4 Character reconstructions

This page provides definitions for each of the characters in our matrix, and justifies codings in particular taxa where relevant. Further citations for codings that are not discussed in the text can be viewed by browsing the morphological dataset on MorphoBank (project 3262).

Alongside its definition, each character has been mapped onto a tree. Any of the optimal trees can be selected by modifying the tree number listed above each diagram. Each tip is labelled according to its coding in the matrix. These states have been used to reconstruct the condition of each internal node, using the parsimony method of Brazeau et al. (2018) as implemented in the R package Inapp.

We emphasize that different trees give different reconstructions. The character mappings are not intended to definitively establish how each character evolved, but to help the reader quickly establish how each character has been coded, and to visualize at a glance how each character fits onto a given tree. Click here to hide the character reconstructions below.

Blue: Logically valid character transformationsOrange: Applicable regionsRed: Logically invalid Fitch stepsOrthrozanclusHalkieria evangelistaSerpulaCapitellaCanadia spinosaSipunculusKulindroplax perissokomosPhthipodochiton thraivensisOdontogriphus omalusWiwaxia corrugataMopaliaTonicellaPolysacos vickersianumLeptochitonGlaphurochiton carbonariusPojetaia runnegariMytilusConocardium elongatumNeopilinaHaliotisPelagiellaDentaliumAcaenoplax hayaeWireniaChaetodermaCalvapilosa kroegeriSiphogonuchites multaLoxosomellaParamicrocornusHaplophrentis carinatusCupitheca holocyclataPedunculotheca dianiaBactrothecaMickwitzia muralensisHeliomedusa orientaMicromitraAskepasma toddenseOrthisTerebratulinaAlisinaGlyptoriaKutorgina chengjiangensisNisusia sulcataAntigonambonites planusCoolinia pectenTomteluva perturbataYuganotheca elegansSalanygolinaLongtancunella chengjiangensisNovocraniaCraniopsUssuniaGasconsiaAcanthotretella spinosaSiphonobolus priscusClupeafumosus socialisLingulosacculusBotsfordiaMummpikia nudaEoobolusLingulellotreta malongensisLingulaPelagodiscus atlanticusMicrinaPaterimitraNamacalathusPhoronisAmathiaFlustraCotyledion tylodesEccentrothecaDailyatiaYilingia spiciformis369413130812736210301610121524162961011271122112115615337111385738115185161186165103219163375344468534228241614421219523211461571054481719116
Click on the numbers by a node for details
Steps incurred at node 74:
25Body organization: Serial repetition168Sclerites: Ornament: Concentric ornament172Sclerites: Composition: Mineralogy
Steps incurred at node 75:
92Digestive tract: Anus: Location96Sclerites: Periodically shed and replaced97Sclerites: Prominent major valves99Sclerites: Accessory sclerites: Arrangement100Sclerites: Accessory sclerites: Symmetry172Sclerites: Composition: Mineralogy176Sclerites: Composition: Microstructure: Format179Sclerites: Structure: Stratiform laminae with polygonal ornament183Sclerites: Structure: External polygonal ornament
114Sclerites: Bivalved: Hinge line shape134Sclerites: Dorsal valve: Growth direction135Sclerites: Dorsal valve: Aspect177Sclerites: Structure: Stratiform lamellae expressed at surface178Sclerites: Structure: Stratiform laminae separated179Sclerites: Structure: Stratiform laminae with polygonal ornament
104Sclerites: Posterior valves: Serially repeated135Sclerites: Dorsal valve: Aspect149Sclerites: Ventral valve: Growth direction150Sclerites: Ventral valve: Relative size
Steps incurred at node 76:
95Sclerites: Present in adult
98Sclerites: Accessory sclerites: Reduced172Sclerites: Composition: Mineralogy175Sclerites: Composition: Microstructure: Number of distinct layers
Steps incurred at node 77:
28Body organization: Pedal groove34Body organization: Circulatory system61Perioral tentacular apparatus: Innervation92Digestive tract: Anus: Location93Digestive tract: Anus: Migration: Within ring of tentacles191Gametes: Gonad position217Embryo: Micromere size219Embryo: Cleavage: Cross pattern221Embryo: Cleavage: Spiral224Larva: Apical organ: Serotonergic cells227Larva: Origin of body cavity229Larva: Foot238Larva: Cilia: Ciliary bands: Downstream246Ciliary ultrastructure: Brushborder of microvilli249Ciliary ultrastructure: Monociliate epidermal cells250Ciliary ultrastructure: Compound cilia: Presence251Ciliary ultrastructure: Compound cilia: Origin253Ciliary ultrastructure: Microvilli on epidermal surface: Branched265Cuticle: Layers266Cuticle: Composition267Cuticle: Fibrous layer with thick fibrils281Nervous system: Buccal nerve ring282Nervous system: Anterior nerve loop291Nervous system: Nerve cords293Nervous system: Ventral longitudinal nerves294Nervous system: Ventral cord location309MicroRNA: mir-1989310MicroRNA: mir-1990311MicroRNA: mir-1991312MicroRNA: mir-1994
94Digestive tract: Anus: Migration: Position
13Adult setae: Secretion: Microvillar canal aspect103Sclerites: Prominent major valves: Additional valves: Nature118Sclerites: Bivalved: Commissure: Circular119Sclerites: Bivalved: Commissure: Lateral margins155Sclerites: Ventral valve: Posterior surface: Extent160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent161Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Identity175Sclerites: Composition: Microstructure: Number of distinct layers
Steps incurred at node 78:
62Perioral tentacular apparatus: Inner nerve ring63Perioral tentacular apparatus: Outer nerve ring85Digestive tract: Oesophageal folds94Digestive tract: Anus: Migration: Position199Gametes: Egg: Site of maturation200Gametes: Spermatozoa: Nucleus: Aspect203Gametes: Spermatozoa: Anterior nuclear fossa205Gametes: Spermatozoa: Acrosome: Shape211Gametes: Spermatozoa: Centrioles: Orientation212Gametes: Spermatozoa: Centrioles: Fusion218Embryo: Cleavage: Equal222Embryo: Origin of mesoderm223Larva: Apical organ: Muscles extending to the hyposphere226Larva: Brain persists into adulthood234Larva: Feeding235Larva: Cilia: Metatroch239Larva: Cilia: Ciliary bands: Upstream240Larva: Cilia: Adoral ciliary band242Ciliary ultrastructure: Accessory centriole257Ciliary ultrastructure: Secondary ciliary rootlet: Length261Nephridia: Serve as excretory organs264Nephridia: Metanephridia268Cuticle: Homogeneous layer273Muscles: Cytology276Nervous system: Orthogonal277Nervous system: Glial system285Nervous system: Formation of ganglia
173Sclerites: Composition: Cuticle or organic matrix
12Adult setae: Secretion: Microvillar diameter17Adult setae: Distribution19Adult setae: Internal constitution
Steps incurred at node 79:
10Adult setae287Nervous system: Cerebral gangila: Fused
11Adult setae: Secretion12Adult setae: Secretion: Microvillar diameter13Adult setae: Secretion: Microvillar canal aspect14Adult setae: Composition: Organic constituent17Adult setae: Distribution19Adult setae: Internal constitution
Steps incurred at node 80:
1Brephic shell: Embryonic shell55Perioral tentacular apparatus: Origin56Perioral tentacular apparatus: Tentacle disposition60Perioral tentacular apparatus: Site of tentacle addition65Perioral tentacular apparatus: Forms closed loop84Digestive tract: Prominent pharynx173Sclerites: Composition: Cuticle or organic matrix174Sclerites: Composition: Incorporation of sedimentary particles180Sclerites: Structure: Canals190Gametes: Gonocoel194Gametes: Asexual reproduction195Gametes: Sexes196Gametes: Fertilization197Gametes: Egg: Size199Gametes: Egg: Site of maturation214Gametes: Spermatozoa: Mitochondria: Shape216Gametes: Spermatozoa: Mitochondria: Midpiece221Embryo: Cleavage: Spiral237Larva: Cilia: Ciliated food groove241Larva: Cilia: Nerve ring underlying ciliated larval swimming organ246Ciliary ultrastructure: Brushborder of microvilli250Ciliary ultrastructure: Compound cilia: Presence259Nephridia: Podocytes263Nephridia: Protonephridia267Cuticle: Fibrous layer with thick fibrils270Cuticle: Microvilli273Muscles: Cytology286Nervous system: Cerebral ganglia: Presence296MicroRNA: Brachiopod candidate 1330MicroRNA: mir-5045
2Brephic shell: Morphology3Brephic shell: Embryonic shell extended in larvae4Brephic shell: Surface ornament5Brephic shell: Larval attachment structure172Sclerites: Composition: Mineralogy175Sclerites: Composition: Microstructure: Number of distinct layers176Sclerites: Composition: Microstructure: Format177Sclerites: Structure: Stratiform lamellae expressed at surface178Sclerites: Structure: Stratiform laminae separated179Sclerites: Structure: Stratiform laminae with polygonal ornament
Steps incurred at node 81:
95Sclerites: Present in adult97Sclerites: Prominent major valves102Sclerites: Prominent major valves: Additional major valves125Sclerites: Bivalved: Muscle scars: Ventral 136Sclerites: Dorsal valve: Posterior surface: Differentiated172Sclerites: Composition: Mineralogy175Sclerites: Composition: Microstructure: Number of distinct layers177Sclerites: Structure: Stratiform lamellae expressed at surface178Sclerites: Structure: Stratiform laminae separated179Sclerites: Structure: Stratiform laminae with polygonal ornament
98Sclerites: Accessory sclerites: Reduced101Sclerites: Prominent major valves: Side slope shape103Sclerites: Prominent major valves: Additional valves: Nature114Sclerites: Bivalved: Hinge line shape116Sclerites: Bivalved: Commissure: Exact correspondence of valve margins118Sclerites: Bivalved: Commissure: Circular126Sclerites: Bivalved: Muscle scars: Ventral: Position128Sclerites: Bivalved: Muscle scars: Dorsal adductors134Sclerites: Dorsal valve: Growth direction135Sclerites: Dorsal valve: Aspect149Sclerites: Ventral valve: Growth direction150Sclerites: Ventral valve: Relative size152Sclerites: Ventral valve: Posterior margin growth direction155Sclerites: Ventral valve: Posterior surface: Extent168Sclerites: Ornament: Concentric ornament169Sclerites: Ornament: Concentric ornament: Symmetry
Steps incurred at node 82:
2Brephic shell: Morphology9Larval setae: Paired bundles10Adult setae15Adult setae: Composition: Enamel35Pedicle: Presence58Perioral tentacular apparatus: Tentacle rows per side in post-trocholophe stage94Digestive tract: Anus: Migration: Position115Sclerites: Bivalved: Enclosing filtration chamber145Sclerites: Dorsal valve: Cardinal processes149Sclerites: Ventral valve: Growth direction151Sclerites: Ventral valve: Posterior surface: Differentiated156Sclerites: Ventral valve: Posterior surface: Delthyrium169Sclerites: Ornament: Concentric ornament: Symmetry172Sclerites: Composition: Mineralogy176Sclerites: Composition: Microstructure: Format
11Adult setae: Secretion12Adult setae: Secretion: Microvillar diameter13Adult setae: Secretion: Microvillar canal aspect14Adult setae: Composition: Organic constituent19Adult setae: Internal constitution37Pedicle: Biomineralization117Sclerites: Bivalved: Commissure: Sulcate157Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape159Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent161Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Identity162Sclerites: Ventral valve: Posterior surface: Delthyrium: Pseudodeltidium: Shape
37Pedicle: Biomineralization165Sclerites: Ventral valve: Umbonal perforation: Shape
Steps incurred at node 83:
2Brephic shell: Morphology4Brephic shell: Surface ornament5Brephic shell: Larval attachment structure6Brephic shell: Setulose7Brephic shell: Setal sacs49Mantle canals: Presence51Mantle canals: vascula lateralia52Mantle canals: vascula media98Sclerites: Accessory sclerites: Reduced100Sclerites: Accessory sclerites: Symmetry116Sclerites: Bivalved: Commissure: Exact correspondence of valve margins134Sclerites: Dorsal valve: Growth direction135Sclerites: Dorsal valve: Aspect137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology180Sclerites: Structure: Canals181Sclerites: Structure: Punctae
8Brephic shell: Setal sacs: Number17Adult setae: Distribution50Mantle canals: Morphology53Mantle canals: vascula terminalia
Steps incurred at node 84:
8Brephic shell: Setal sacs: Number115Sclerites: Bivalved: Enclosing filtration chamber120Sclerites: Bivalved: Apophyses136Sclerites: Dorsal valve: Posterior surface: Differentiated155Sclerites: Ventral valve: Posterior surface: Extent160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent177Sclerites: Structure: Stratiform lamellae expressed at surface178Sclerites: Structure: Stratiform laminae separated179Sclerites: Structure: Stratiform laminae with polygonal ornament
117Sclerites: Bivalved: Commissure: Sulcate121Sclerites: Bivalved: Apophyses: Morphology
Steps incurred at node 85:
2Brephic shell: Morphology17Adult setae: Distribution65Perioral tentacular apparatus: Forms closed loop66Perioral tentacular apparatus: Coiling direction128Sclerites: Bivalved: Muscle scars: Dorsal adductors132Sclerites: Bivalved: Muscle scars: Dorsal diductor134Sclerites: Dorsal valve: Growth direction169Sclerites: Ornament: Concentric ornament: Symmetry180Sclerites: Structure: Canals181Sclerites: Structure: Punctae
36Pedicle: Constitution40Pedicle: Tapering41Pedicle: Coelomic region42Pedicle: Surface ornament129Sclerites: Bivalved: Muscle scars: Adductors: Position133Sclerites: Bivalved: Muscle scars: Dorsal diductor: Position
158Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape: Aspect of rounded opening
Steps incurred at node 86:
8Brephic shell: Setal sacs: Number23Adult setae: Projecting knobs40Pedicle: Tapering42Pedicle: Surface ornament59Perioral tentacular apparatus: Median tentacle in early development67Perioral tentacular apparatus: Adjustor muscle114Sclerites: Bivalved: Hinge line shape118Sclerites: Bivalved: Commissure: Circular136Sclerites: Dorsal valve: Posterior surface: Differentiated137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology142Sclerites: Dorsal valve: Notothyrial platform150Sclerites: Ventral valve: Relative size153Sclerites: Ventral valve: Posterior surface: Planar162Sclerites: Ventral valve: Posterior surface: Delthyrium: Pseudodeltidium: Shape167Sclerites: Ventral valve: Median septum170Sclerites: Ornament: Radial ornament203Gametes: Spermatozoa: Anterior nuclear fossa205Gametes: Spermatozoa: Acrosome: Shape206Gametes: Spermatozoa: Acrosome: Differentiated internally209Gametes: Spermatozoa: Mid-piece211Gametes: Spermatozoa: Centrioles: Orientation224Larva: Apical organ: Serotonergic cells235Larva: Cilia: Metatroch255Ciliary ultrastructure: Vertical ciliary rootlet: Shape257Ciliary ultrastructure: Secondary ciliary rootlet: Length268Cuticle: Homogeneous layer276Nervous system: Orthogonal
119Sclerites: Bivalved: Commissure: Lateral margins
119Sclerites: Bivalved: Commissure: Lateral margins
Steps incurred at node 87:
5Brephic shell: Larval attachment structure10Adult setae15Adult setae: Composition: Enamel35Pedicle: Presence43Pedicle: Nerve impression50Mantle canals: Morphology53Mantle canals: vascula terminalia57Perioral tentacular apparatus: Tentacle rows per side in trocholophe stage92Digestive tract: Anus: Location128Sclerites: Bivalved: Muscle scars: Dorsal adductors130Sclerites: Bivalved: Muscle scars: Dermal muscles132Sclerites: Bivalved: Muscle scars: Dorsal diductor135Sclerites: Dorsal valve: Aspect157Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape168Sclerites: Ornament: Concentric ornament172Sclerites: Composition: Mineralogy173Sclerites: Composition: Cuticle or organic matrix197Gametes: Egg: Size199Gametes: Egg: Site of maturation205Gametes: Spermatozoa: Acrosome: Shape224Larva: Apical organ: Serotonergic cells279Nervous system: Statocysts
158Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape: Aspect of rounded opening
160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent
Steps incurred at node 88:
8Brephic shell: Setal sacs: Number23Adult setae: Projecting knobs24Adult setae: Circling coronae38Pedicle: Bulb65Perioral tentacular apparatus: Forms closed loop67Perioral tentacular apparatus: Adjustor muscle84Digestive tract: Prominent pharynx93Digestive tract: Anus: Migration: Within ring of tentacles94Digestive tract: Anus: Migration: Position118Sclerites: Bivalved: Commissure: Circular128Sclerites: Bivalved: Muscle scars: Dorsal adductors134Sclerites: Dorsal valve: Growth direction135Sclerites: Dorsal valve: Aspect136Sclerites: Dorsal valve: Posterior surface: Differentiated138Sclerites: Dorsal valve: Posterior surface: Medial groove143Sclerites: Dorsal valve: Medial septum149Sclerites: Ventral valve: Growth direction150Sclerites: Ventral valve: Relative size151Sclerites: Ventral valve: Posterior surface: Differentiated180Sclerites: Structure: Canals276Nervous system: Orthogonal
119Sclerites: Bivalved: Commissure: Lateral margins
Steps incurred at node 89:
3Brephic shell: Embryonic shell extended in larvae7Brephic shell: Setal sacs40Pedicle: Tapering42Pedicle: Surface ornament50Mantle canals: Morphology52Mantle canals: vascula media58Perioral tentacular apparatus: Tentacle rows per side in post-trocholophe stage119Sclerites: Bivalved: Commissure: Lateral margins134Sclerites: Dorsal valve: Growth direction149Sclerites: Ventral valve: Growth direction150Sclerites: Ventral valve: Relative size155Sclerites: Ventral valve: Posterior surface: Extent156Sclerites: Ventral valve: Posterior surface: Delthyrium167Sclerites: Ventral valve: Median septum181Sclerites: Structure: Punctae
Steps incurred at node 90:
17Adult setae: Distribution43Pedicle: Nerve impression118Sclerites: Bivalved: Commissure: Circular152Sclerites: Ventral valve: Posterior margin growth direction153Sclerites: Ventral valve: Posterior surface: Planar158Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape: Aspect of rounded opening
Steps incurred at node 91:
3Brephic shell: Embryonic shell extended in larvae4Brephic shell: Surface ornament150Sclerites: Ventral valve: Relative size155Sclerites: Ventral valve: Posterior surface: Extent157Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape
159Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover
Steps incurred at node 92:
8Brephic shell: Setal sacs: Number137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology159Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover
160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent161Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Identity162Sclerites: Ventral valve: Posterior surface: Delthyrium: Pseudodeltidium: Shape
Steps incurred at node 93:
143Sclerites: Dorsal valve: Medial septum150Sclerites: Ventral valve: Relative size153Sclerites: Ventral valve: Posterior surface: Planar157Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape172Sclerites: Composition: Mineralogy175Sclerites: Composition: Microstructure: Number of distinct layers180Sclerites: Structure: Canals
Steps incurred at node 94:
4Brephic shell: Surface ornament5Brephic shell: Larval attachment structure50Mantle canals: Morphology134Sclerites: Dorsal valve: Growth direction137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology171Sclerites: Ornament: Shell-penetrating spines172Sclerites: Composition: Mineralogy178Sclerites: Structure: Stratiform laminae separated179Sclerites: Structure: Stratiform laminae with polygonal ornament180Sclerites: Structure: Canals181Sclerites: Structure: Punctae
165Sclerites: Ventral valve: Umbonal perforation: Shape
Steps incurred at node 95:
155Sclerites: Ventral valve: Posterior surface: Extent166Sclerites: Ventral valve: Colleplax, cicatrix or pedicle sheath170Sclerites: Ornament: Radial ornament
Steps incurred at node 96:
114Sclerites: Bivalved: Hinge line shape123Sclerites: Bivalved: Sockets131Sclerites: Bivalved: Muscle scars: Unpaired median (levator ani)151Sclerites: Ventral valve: Posterior surface: Differentiated153Sclerites: Ventral valve: Posterior surface: Planar156Sclerites: Ventral valve: Posterior surface: Delthyrium167Sclerites: Ventral valve: Median septum170Sclerites: Ornament: Radial ornament
Steps incurred at node 97:
128Sclerites: Bivalved: Muscle scars: Dorsal adductors134Sclerites: Dorsal valve: Growth direction149Sclerites: Ventral valve: Growth direction168Sclerites: Ornament: Concentric ornament172Sclerites: Composition: Mineralogy
Steps incurred at node 98:
1Brephic shell: Embryonic shell114Sclerites: Bivalved: Hinge line shape136Sclerites: Dorsal valve: Posterior surface: Differentiated143Sclerites: Dorsal valve: Medial septum150Sclerites: Ventral valve: Relative size166Sclerites: Ventral valve: Colleplax, cicatrix or pedicle sheath181Sclerites: Structure: Punctae
Steps incurred at node 99:
1Brephic shell: Embryonic shell4Brephic shell: Surface ornament120Sclerites: Bivalved: Apophyses139Sclerites: Dorsal valve: Posterior surface: Notothyrium168Sclerites: Ornament: Concentric ornament172Sclerites: Composition: Mineralogy175Sclerites: Composition: Microstructure: Number of distinct layers176Sclerites: Composition: Microstructure: Format
121Sclerites: Bivalved: Apophyses: Morphology140Sclerites: Dorsal valve: Posterior surface: Notothyrium: Shape141Sclerites: Dorsal valve: Posterior surface: Notothyrium: Chilidial plates
Steps incurred at node 100:
39Pedicle: Distal rootlets41Pedicle: Coelomic region56Perioral tentacular apparatus: Tentacle disposition58Perioral tentacular apparatus: Tentacle rows per side in post-trocholophe stage91Digestive tract: Anus: Presence94Digestive tract: Anus: Migration: Position127Sclerites: Bivalved: Muscle scars: Adjustor133Sclerites: Bivalved: Muscle scars: Dorsal diductor: Position141Sclerites: Dorsal valve: Posterior surface: Notothyrium: Chilidial plates142Sclerites: Dorsal valve: Notothyrial platform150Sclerites: Ventral valve: Relative size159Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover161Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Identity163Sclerites: Ventral valve: Posterior surface: Delthyrium: Pseudodeltidium: Hinge furrows164Sclerites: Ventral valve: Umbonal perforation
165Sclerites: Ventral valve: Umbonal perforation: Shape
162Sclerites: Ventral valve: Posterior surface: Delthyrium: Pseudodeltidium: Shape
Steps incurred at node 101:
1Brephic shell: Embryonic shell65Perioral tentacular apparatus: Forms closed loop92Digestive tract: Anus: Location94Digestive tract: Anus: Migration: Position121Sclerites: Bivalved: Apophyses: Morphology166Sclerites: Ventral valve: Colleplax, cicatrix or pedicle sheath170Sclerites: Ornament: Radial ornament176Sclerites: Composition: Microstructure: Format
Steps incurred at node 102:
36Pedicle: Constitution37Pedicle: Biomineralization41Pedicle: Coelomic region52Mantle canals: vascula media119Sclerites: Bivalved: Commissure: Lateral margins
Steps incurred at node 103:
2Brephic shell: Morphology38Pedicle: Bulb120Sclerites: Bivalved: Apophyses122Sclerites: Bivalved: Apophyses: Dental plates123Sclerites: Bivalved: Sockets124Sclerites: Bivalved: Socket ridges142Sclerites: Dorsal valve: Notothyrial platform145Sclerites: Dorsal valve: Cardinal processes150Sclerites: Ventral valve: Relative size153Sclerites: Ventral valve: Posterior surface: Planar157Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape170Sclerites: Ornament: Radial ornament177Sclerites: Structure: Stratiform lamellae expressed at surface178Sclerites: Structure: Stratiform laminae separated179Sclerites: Structure: Stratiform laminae with polygonal ornament182Sclerites: Structure: Pseudopunctae
169Sclerites: Ornament: Concentric ornament: Symmetry
Steps incurred at node 104:
1Brephic shell: Embryonic shell114Sclerites: Bivalved: Hinge line shape119Sclerites: Bivalved: Commissure: Lateral margins137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology149Sclerites: Ventral valve: Growth direction160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent161Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Identity168Sclerites: Ornament: Concentric ornament
169Sclerites: Ornament: Concentric ornament: Symmetry
Steps incurred at node 105:
117Sclerites: Bivalved: Commissure: Sulcate118Sclerites: Bivalved: Commissure: Circular145Sclerites: Dorsal valve: Cardinal processes170Sclerites: Ornament: Radial ornament172Sclerites: Composition: Mineralogy174Sclerites: Composition: Incorporation of sedimentary particles
Steps incurred at node 106:
134Sclerites: Dorsal valve: Growth direction143Sclerites: Dorsal valve: Medial septum165Sclerites: Ventral valve: Umbonal perforation: Shape167Sclerites: Ventral valve: Median septum168Sclerites: Ornament: Concentric ornament175Sclerites: Composition: Microstructure: Number of distinct layers
169Sclerites: Ornament: Concentric ornament: Symmetry
Steps incurred at node 107:
36Pedicle: Constitution40Pedicle: Tapering141Sclerites: Dorsal valve: Posterior surface: Notothyrium: Chilidial plates160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent171Sclerites: Ornament: Shell-penetrating spines
Steps incurred at node 108:
117Sclerites: Bivalved: Commissure: Sulcate118Sclerites: Bivalved: Commissure: Circular119Sclerites: Bivalved: Commissure: Lateral margins123Sclerites: Bivalved: Sockets124Sclerites: Bivalved: Socket ridges126Sclerites: Bivalved: Muscle scars: Ventral: Position128Sclerites: Bivalved: Muscle scars: Dorsal adductors129Sclerites: Bivalved: Muscle scars: Adductors: Position133Sclerites: Bivalved: Muscle scars: Dorsal diductor: Position145Sclerites: Dorsal valve: Cardinal processes
Steps incurred at node 109:
36Pedicle: Constitution38Pedicle: Bulb40Pedicle: Tapering42Pedicle: Surface ornament50Mantle canals: Morphology114Sclerites: Bivalved: Hinge line shape118Sclerites: Bivalved: Commissure: Circular121Sclerites: Bivalved: Apophyses: Morphology122Sclerites: Bivalved: Apophyses: Dental plates137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology139Sclerites: Dorsal valve: Posterior surface: Notothyrium140Sclerites: Dorsal valve: Posterior surface: Notothyrium: Shape142Sclerites: Dorsal valve: Notothyrial platform143Sclerites: Dorsal valve: Medial septum145Sclerites: Dorsal valve: Cardinal processes159Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent164Sclerites: Ventral valve: Umbonal perforation168Sclerites: Ornament: Concentric ornament170Sclerites: Ornament: Radial ornament181Sclerites: Structure: Punctae
162Sclerites: Ventral valve: Posterior surface: Delthyrium: Pseudodeltidium: Shape165Sclerites: Ventral valve: Umbonal perforation: Shape169Sclerites: Ornament: Concentric ornament: Symmetry
Steps incurred at node 110:
4Brephic shell: Surface ornament50Mantle canals: Morphology126Sclerites: Bivalved: Muscle scars: Ventral: Position159Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover167Sclerites: Ventral valve: Median septum169Sclerites: Ornament: Concentric ornament: Symmetry170Sclerites: Ornament: Radial ornament177Sclerites: Structure: Stratiform lamellae expressed at surface179Sclerites: Structure: Stratiform laminae with polygonal ornament
Steps incurred at node 111:
35Pedicle: Presence125Sclerites: Bivalved: Muscle scars: Ventral 143Sclerites: Dorsal valve: Medial septum149Sclerites: Ventral valve: Growth direction156Sclerites: Ventral valve: Posterior surface: Delthyrium170Sclerites: Ornament: Radial ornament
158Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape: Aspect of rounded opening
Steps incurred at node 112:
144Sclerites: Dorsal valve: Cardinal shield153Sclerites: Ventral valve: Posterior surface: Planar168Sclerites: Ornament: Concentric ornament
Steps incurred at node 113:
2Brephic shell: Morphology116Sclerites: Bivalved: Commissure: Exact correspondence of valve margins118Sclerites: Bivalved: Commissure: Circular147Sclerites: Dorsal valve: Clavicles151Sclerites: Ventral valve: Posterior surface: Differentiated154Sclerites: Ventral valve: Ligula164Sclerites: Ventral valve: Umbonal perforation
119Sclerites: Bivalved: Commissure: Lateral margins148Sclerites: Dorsal valve: Clavicles: Type of clavicles165Sclerites: Ventral valve: Umbonal perforation: Shape
Steps incurred at node 114:
35Pedicle: Presence39Pedicle: Distal rootlets153Sclerites: Ventral valve: Posterior surface: Planar
36Pedicle: Constitution37Pedicle: Biomineralization40Pedicle: Tapering41Pedicle: Coelomic region42Pedicle: Surface ornament
Steps incurred at node 115:
98Sclerites: Accessory sclerites: Reduced100Sclerites: Accessory sclerites: Symmetry137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology168Sclerites: Ornament: Concentric ornament
Steps incurred at node 116:
10Adult setae25Body organization: Serial repetition29Body organization: Pedal groove: Foot30Body organization: Coelom31Body organization: Coelomoducts: Number54Perioral tentacular apparatus: Presence55Perioral tentacular apparatus: Origin64Perioral tentacular apparatus: Musculature92Digestive tract: Anus: Location190Gametes: Gonocoel194Gametes: Asexual reproduction195Gametes: Sexes196Gametes: Fertilization197Gametes: Egg: Size198Gametes: Egg: Protective membrane203Gametes: Spermatozoa: Anterior nuclear fossa204Gametes: Spermatozoa: Acrosome206Gametes: Spermatozoa: Acrosome: Differentiated internally209Gametes: Spermatozoa: Mid-piece210Gametes: Spermatozoa: Mid-piece: Mitochondrial location214Gametes: Spermatozoa: Mitochondria: Shape215Gametes: Spermatozoa: Mitochondria: Cristae: Configuration216Gametes: Spermatozoa: Mitochondria: Midpiece220Embryo: Cleavage: Polar lobe formation225Larva: Apical organ: Develops into adult brain230Larva: Foot: Pedal gland231Larva: Foot: Pedal gland: Retained to adulthood232Larva: Coelom: Paired234Larva: Feeding237Larva: Cilia: Ciliated food groove243Ciliary ultrastructure: Aggregation of granules below basal plate244Ciliary ultrastructure: Basal foot: Radiating tubular fibres245Ciliary ultrastructure: Basal plate247Ciliary ultrastructure: Centriolar triplet derivative in basal body248Ciliary ultrastructure: Ciliary necklace with connecting strands254Ciliary ultrastructure: Vertical ciliary rootlet: Length255Ciliary ultrastructure: Vertical ciliary rootlet: Shape256Ciliary ultrastructure: Secondary ciliary rootlet: Presence259Nephridia: Podocytes261Nephridia: Serve as excretory organs263Nephridia: Protonephridia269Cuticle: Resilience270Cuticle: Microvilli277Nervous system: Glial system283Nervous system: Suprarectal commissure285Nervous system: Formation of ganglia
11Adult setae: Secretion14Adult setae: Composition: Organic constituent17Adult setae: Distribution19Adult setae: Internal constitution
Steps incurred at node 117:
16Adult setae: Composition: Mineralized core17Adult setae: Distribution26Body organization: Annulae27Body organization: Subdivided head28Body organization: Pedal groove32Body organization: Gills34Body organization: Circulatory system44Mantle cavity: Presence45Mantle cavity: Reduced68Mouthparts: Buccal organ84Digestive tract: Prominent pharynx85Digestive tract: Oesophageal folds86Digestive tract: Oral sphincter88Digestive tract: Foregut: Locomotory cilia89Digestive tract: Midgut: Subdivisions90Digestive tract: Midgut: Glands95Sclerites: Present in adult97Sclerites: Prominent major valves191Gametes: Gonad position192Gametes: Ovary wall saccular193Gametes: Testis wall saccular199Gametes: Egg: Site of maturation200Gametes: Spermatozoa: Nucleus: Aspect205Gametes: Spermatozoa: Acrosome: Shape207Gametes: Spermatozoa: Acrosome: Subacrosomal basal plate210Gametes: Spermatozoa: Mid-piece: Mitochondrial location213Gametes: Spermatozoa: Satellite fibre complex219Embryo: Cleavage: Cross pattern228Larva: Formation of coelomoducts233Larva: Coelom: Paried: Includes pericardium235Larva: Cilia: Metatroch236Larva: Cilia: Telotroch260Nephridia: Rhogocytes264Nephridia: Metanephridia266Cuticle: Composition267Cuticle: Fibrous layer with thick fibrils268Cuticle: Homogeneous layer272Muscles: Circular muscles273Muscles: Cytology274Muscles: Histology275Glands: Paired pharyngeal diverticulae278Nervous system: Dorso-terminal sense organ280Nervous system: Nuchal organs288Nervous system: Cerebral ganglia: Transverse commissures291Nervous system: Nerve cords313MicroRNA: mir-1995314MicroRNA: mir-1996315MicroRNA: mir-1997318MicroRNA: mir-2000320MicroRNA: mir-2685324MicroRNA: mir-2689326MicroRNA: mir-2691329MicroRNA: mir-2722
33Body organization: Gills: Ctenidia46Mantle cavity: Open face47Mantle cavity: Sac opening69Mouthparts: Buccal organ: Eversible98Sclerites: Accessory sclerites: Reduced101Sclerites: Prominent major valves: Side slope shape135Sclerites: Dorsal valve: Aspect168Sclerites: Ornament: Concentric ornament
12Adult setae: Secretion: Microvillar diameter55Perioral tentacular apparatus: Origin64Perioral tentacular apparatus: Musculature65Perioral tentacular apparatus: Forms closed loop
Steps incurred at node 118:
10Adult setae11Adult setae: Secretion16Adult setae: Composition: Mineralized core17Adult setae: Distribution19Adult setae: Internal constitution20Adult setae: Distinct shaft25Body organization: Serial repetition28Body organization: Pedal groove29Body organization: Pedal groove: Foot33Body organization: Gills: Ctenidia45Mantle cavity: Reduced46Mantle cavity: Open face71Radula72Radula: Extent73Radula: Subradular membrane74Radula: Alary processes75Radula: Bolster vesicles76Radula: Subradular organ77Radula: Teeth: Heterodonty79Radula: Teeth: More teeth per row in larger individuals80Radula: Teeth: Lateral tooth base81Radula: Teeth: Lateral tooth head82Radula: Teeth: Apatite83Radula: Teeth: Magnetite89Digestive tract: Midgut: Subdivisions90Digestive tract: Midgut: Glands92Digestive tract: Anus: Location95Sclerites: Present in adult97Sclerites: Prominent major valves102Sclerites: Prominent major valves: Additional major valves106Sclerites: Posterior valves: Apophyses112Sclerites: Posterior valves: Differentiated intermediate shell fields134Sclerites: Dorsal valve: Growth direction135Sclerites: Dorsal valve: Aspect136Sclerites: Dorsal valve: Posterior surface: Differentiated137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology168Sclerites: Ornament: Concentric ornament184Sclerites: Structure: Aesthete canals189Sclerites: Structure: Articulamentum195Gametes: Sexes196Gametes: Fertilization198Gametes: Egg: Protective membrane200Gametes: Spermatozoa: Nucleus: Aspect201Gametes: Spermatozoa: Nucleus: Shape205Gametes: Spermatozoa: Acrosome: Shape206Gametes: Spermatozoa: Acrosome: Differentiated internally207Gametes: Spermatozoa: Acrosome: Subacrosomal basal plate208Gametes: Spermatozoa: Acrosome: Subacrosomal basal plate: Basal granule210Gametes: Spermatozoa: Mid-piece: Mitochondrial location214Gametes: Spermatozoa: Mitochondria: Shape216Gametes: Spermatozoa: Mitochondria: Midpiece217Embryo: Micromere size218Embryo: Cleavage: Equal219Embryo: Cleavage: Cross pattern220Embryo: Cleavage: Polar lobe formation223Larva: Apical organ: Muscles extending to the hyposphere224Larva: Apical organ: Serotonergic cells230Larva: Foot: Pedal gland231Larva: Foot: Pedal gland: Retained to adulthood236Larva: Cilia: Telotroch246Ciliary ultrastructure: Brushborder of microvilli252Ciliary ultrastructure: Glycocalyx ultrastructure258Ciliary ultrastructure: Secondary ciliary rootlet: Shape269Cuticle: Resilience271Muscles: Longitudinal muscle bands278Nervous system: Dorso-terminal sense organ279Nervous system: Statocysts281Nervous system: Buccal nerve ring283Nervous system: Suprarectal commissure284Nervous system: Suprarectal commissure: Ganglionated285Nervous system: Formation of ganglia287Nervous system: Cerebral gangila: Fused297MicroRNA: mir-36299MicroRNA: mir-124300MicroRNA: mir-190301MicroRNA: mir-219302MicroRNA: mir-242304MicroRNA: mir-1984305MicroRNA: mir-1985306MicroRNA: mir-1986308MicroRNA: mir-1988309MicroRNA: mir-1989
12Adult setae: Secretion: Microvillar diameter13Adult setae: Secretion: Microvillar canal aspect18Adult setae: Distribution: Present on anteriormost segment47Mantle cavity: Sac opening72Radula: Extent77Radula: Teeth: Heterodonty78Radula: Teeth: Bending plane79Radula: Teeth: More teeth per row in larger individuals103Sclerites: Prominent major valves: Additional valves: Nature104Sclerites: Posterior valves: Serially repeated105Sclerites: Posterior valves: Number112Sclerites: Posterior valves: Differentiated intermediate shell fields113Sclerites: Posterior valves: Laterally divided shell fields114Sclerites: Bivalved: Hinge line shape134Sclerites: Dorsal valve: Growth direction169Sclerites: Ornament: Concentric ornament: Symmetry172Sclerites: Composition: Mineralogy175Sclerites: Composition: Microstructure: Number of distinct layers176Sclerites: Composition: Microstructure: Format185Sclerites: Structure: Aesthete canals: Orientation186Sclerites: Structure: Aesthete canals: Size variation187Sclerites: Structure: Aesthete canals: Megalaesthete bulbs
47Mantle cavity: Sac opening134Sclerites: Dorsal valve: Growth direction169Sclerites: Ornament: Concentric ornament: Symmetry176Sclerites: Composition: Microstructure: Format
Steps incurred at node 119:
1Brephic shell: Embryonic shell33Body organization: Gills: Ctenidia54Perioral tentacular apparatus: Presence74Radula: Alary processes87Digestive tract: Oral sphincter: Reduced92Digestive tract: Anus: Location170Sclerites: Ornament: Radial ornament206Gametes: Spermatozoa: Acrosome: Differentiated internally212Gametes: Spermatozoa: Centrioles: Fusion236Larva: Cilia: Telotroch256Ciliary ultrastructure: Secondary ciliary rootlet: Presence276Nervous system: Orthogonal292Nervous system: Nerve cords: Medulla293Nervous system: Ventral longitudinal nerves
2Brephic shell: Morphology3Brephic shell: Embryonic shell extended in larvae4Brephic shell: Surface ornament5Brephic shell: Larval attachment structure55Perioral tentacular apparatus: Origin56Perioral tentacular apparatus: Tentacle disposition61Perioral tentacular apparatus: Innervation64Perioral tentacular apparatus: Musculature65Perioral tentacular apparatus: Forms closed loop118Sclerites: Bivalved: Commissure: Circular119Sclerites: Bivalved: Commissure: Lateral margins134Sclerites: Dorsal valve: Growth direction172Sclerites: Composition: Mineralogy173Sclerites: Composition: Cuticle or organic matrix175Sclerites: Composition: Microstructure: Number of distinct layers176Sclerites: Composition: Microstructure: Format
Steps incurred at node 120:
2Brephic shell: Morphology10Adult setae134Sclerites: Dorsal valve: Growth direction176Sclerites: Composition: Microstructure: Format
Steps incurred at node 121:
46Mantle cavity: Open face48Mantle cavity: Pallial line78Radula: Teeth: Bending plane168Sclerites: Ornament: Concentric ornament175Sclerites: Composition: Microstructure: Number of distinct layers196Gametes: Fertilization203Gametes: Spermatozoa: Anterior nuclear fossa210Gametes: Spermatozoa: Mid-piece: Mitochondrial location218Embryo: Cleavage: Equal219Embryo: Cleavage: Cross pattern224Larva: Apical organ: Serotonergic cells287Nervous system: Cerebral gangila: Fused
47Mantle cavity: Sac opening169Sclerites: Ornament: Concentric ornament: Symmetry
Steps incurred at node 122:
2Brephic shell: Morphology25Body organization: Serial repetition71Radula74Radula: Alary processes75Radula: Bolster vesicles76Radula: Subradular organ80Radula: Teeth: Lateral tooth base85Digestive tract: Oesophageal folds87Digestive tract: Oral sphincter: Reduced92Digestive tract: Anus: Location134Sclerites: Dorsal valve: Growth direction138Sclerites: Dorsal valve: Posterior surface: Medial groove191Gametes: Gonad position192Gametes: Ovary wall saccular193Gametes: Testis wall saccular197Gametes: Egg: Size200Gametes: Spermatozoa: Nucleus: Aspect206Gametes: Spermatozoa: Acrosome: Differentiated internally207Gametes: Spermatozoa: Acrosome: Subacrosomal basal plate
Steps incurred at node 123:
136Sclerites: Dorsal valve: Posterior surface: Differentiated137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology168Sclerites: Ornament: Concentric ornament170Sclerites: Ornament: Radial ornament175Sclerites: Composition: Microstructure: Number of distinct layers
Steps incurred at node 124:
134Sclerites: Dorsal valve: Growth direction175Sclerites: Composition: Microstructure: Number of distinct layers
Steps incurred at node 125:
107Sclerites: Posterior valves: Jugal ridges151Sclerites: Ventral valve: Posterior surface: Differentiated
149Sclerites: Ventral valve: Growth direction150Sclerites: Ventral valve: Relative size152Sclerites: Ventral valve: Posterior margin growth direction
Steps incurred at node 126:
14Adult setae: Composition: Organic constituent16Adult setae: Composition: Mineralized core108Sclerites: Posterior valves: Insertion plates112Sclerites: Posterior valves: Differentiated intermediate shell fields150Sclerites: Ventral valve: Relative size168Sclerites: Ornament: Concentric ornament186Sclerites: Structure: Aesthete canals: Size variation202Gametes: Spermatozoa: Nucleus: Nuclear filament205Gametes: Spermatozoa: Acrosome: Shape206Gametes: Spermatozoa: Acrosome: Differentiated internally208Gametes: Spermatozoa: Acrosome: Subacrosomal basal plate: Basal granule210Gametes: Spermatozoa: Mid-piece: Mitochondrial location212Gametes: Spermatozoa: Centrioles: Fusion214Gametes: Spermatozoa: Mitochondria: Shape
111Sclerites: Posterior valves: Insertion plates: Pectinate
Steps incurred at node 127:
98Sclerites: Accessory sclerites: Reduced99Sclerites: Accessory sclerites: Arrangement100Sclerites: Accessory sclerites: Symmetry105Sclerites: Posterior valves: Number109Sclerites: Posterior valves: Insertion plates: Slit111Sclerites: Posterior valves: Insertion plates: Pectinate113Sclerites: Posterior valves: Laterally divided shell fields134Sclerites: Dorsal valve: Growth direction136Sclerites: Dorsal valve: Posterior surface: Differentiated137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology151Sclerites: Ventral valve: Posterior surface: Differentiated156Sclerites: Ventral valve: Posterior surface: Delthyrium170Sclerites: Ornament: Radial ornament172Sclerites: Composition: Mineralogy187Sclerites: Structure: Aesthete canals: Megalaesthete bulbs
110Sclerites: Posterior valves: Insertion plates: Slit: Nature155Sclerites: Ventral valve: Posterior surface: Extent157Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape159Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover160Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent161Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Identity
Steps incurred at node 128:
20Adult setae: Distinct shaft107Sclerites: Posterior valves: Jugal ridges112Sclerites: Posterior valves: Differentiated intermediate shell fields120Sclerites: Bivalved: Apophyses134Sclerites: Dorsal valve: Growth direction137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology139Sclerites: Dorsal valve: Posterior surface: Notothyrium186Sclerites: Structure: Aesthete canals: Size variation200Gametes: Spermatozoa: Nucleus: Aspect207Gametes: Spermatozoa: Acrosome: Subacrosomal basal plate
116Sclerites: Bivalved: Commissure: Exact correspondence of valve margins117Sclerites: Bivalved: Commissure: Sulcate118Sclerites: Bivalved: Commissure: Circular119Sclerites: Bivalved: Commissure: Lateral margins121Sclerites: Bivalved: Apophyses: Morphology140Sclerites: Dorsal valve: Posterior surface: Notothyrium: Shape141Sclerites: Dorsal valve: Posterior surface: Notothyrium: Chilidial plates
Steps incurred at node 129:
Steps incurred at node 130:
28Body organization: Pedal groove47Mantle cavity: Sac opening105Sclerites: Posterior valves: Number136Sclerites: Dorsal valve: Posterior surface: Differentiated137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology
Steps incurred at node 131:
9Larval setae: Paired bundles11Adult setae: Secretion14Adult setae: Composition: Organic constituent19Adult setae: Internal constitution23Adult setae: Projecting knobs31Body organization: Coelomoducts: Number34Body organization: Circulatory system54Perioral tentacular apparatus: Presence56Perioral tentacular apparatus: Tentacle disposition70Mouthparts: Buccal organ: Papillae92Digestive tract: Anus: Location203Gametes: Spermatozoa: Anterior nuclear fossa212Gametes: Spermatozoa: Centrioles: Fusion215Gametes: Spermatozoa: Mitochondria: Cristae: Configuration224Larva: Apical organ: Serotonergic cells227Larva: Origin of body cavity238Larva: Cilia: Ciliary bands: Downstream242Ciliary ultrastructure: Accessory centriole243Ciliary ultrastructure: Aggregation of granules below basal plate246Ciliary ultrastructure: Brushborder of microvilli247Ciliary ultrastructure: Centriolar triplet derivative in basal body248Ciliary ultrastructure: Ciliary necklace with connecting strands252Ciliary ultrastructure: Glycocalyx ultrastructure253Ciliary ultrastructure: Microvilli on epidermal surface: Branched254Ciliary ultrastructure: Vertical ciliary rootlet: Length257Ciliary ultrastructure: Secondary ciliary rootlet: Length262Nephridia: Serially repeated263Nephridia: Protonephridia270Cuticle: Microvilli289Nervous system: Serially repeated ganglia290Nervous system: Serially repeated segmental nerves295Nervous system: Ventral cord commissures297MicroRNA: mir-36298MicroRNA: mir-76299MicroRNA: mir-124302MicroRNA: mir-242303MicroRNA: mir-278307MicroRNA: mir-1987310MicroRNA: mir-1990311MicroRNA: mir-1991316MicroRNA: mir-1998317MicroRNA: mir-1999321MicroRNA: mir-2686322MicroRNA: mir-2687323MicroRNA: mir-2688325MicroRNA: mir-2690327MicroRNA: mir-2693328MicroRNA: mir-2693
12Adult setae: Secretion: Microvillar diameter13Adult setae: Secretion: Microvillar canal aspect18Adult setae: Distribution: Present on anteriormost segment94Digestive tract: Anus: Migration: Position
Steps incurred at node 132:
18Adult setae: Distribution: Present on anteriormost segment20Adult setae: Distinct shaft21Adult setae: Hooks22Adult setae: Capitium26Body organization: Annulae27Body organization: Subdivided head32Body organization: Gills
33Body organization: Gills: Ctenidia
Steps incurred at node 133:
15Adult setae: Composition: Enamel23Adult setae: Projecting knobs34Body organization: Circulatory system54Perioral tentacular apparatus: Presence68Mouthparts: Buccal organ70Mouthparts: Buccal organ: Papillae84Digestive tract: Prominent pharynx196Gametes: Fertilization197Gametes: Egg: Size200Gametes: Spermatozoa: Nucleus: Aspect205Gametes: Spermatozoa: Acrosome: Shape206Gametes: Spermatozoa: Acrosome: Differentiated internally209Gametes: Spermatozoa: Mid-piece211Gametes: Spermatozoa: Centrioles: Orientation214Gametes: Spermatozoa: Mitochondria: Shape216Gametes: Spermatozoa: Mitochondria: Midpiece235Larva: Cilia: Metatroch236Larva: Cilia: Telotroch237Larva: Cilia: Ciliated food groove
Steps incurred at node 134:
102Sclerites: Prominent major valves: Additional major valves134Sclerites: Dorsal valve: Growth direction136Sclerites: Dorsal valve: Posterior surface: Differentiated137Sclerites: Dorsal valve: Differentiated posterior surface: Morphology151Sclerites: Ventral valve: Posterior surface: Differentiated153Sclerites: Ventral valve: Posterior surface: Planar
103Sclerites: Prominent major valves: Additional valves: Nature104Sclerites: Posterior valves: Serially repeated113Sclerites: Posterior valves: Laterally divided shell fields116Sclerites: Bivalved: Commissure: Exact correspondence of valve margins117Sclerites: Bivalved: Commissure: Sulcate118Sclerites: Bivalved: Commissure: Circular119Sclerites: Bivalved: Commissure: Lateral margins149Sclerites: Ventral valve: Growth direction150Sclerites: Ventral valve: Relative size152Sclerites: Ventral valve: Posterior margin growth direction155Sclerites: Ventral valve: Posterior surface: Extent

4.1 Brephic shell

[1] Embryonic shell

Character adds 7 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[0]Dentalium[1]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[1]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[?]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[1]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 1: Brephic shell: Embryonic shell

0: Absent1: Present?: AmbiguousNeomorphic character.

The embryonic shell or protegulum is secreted by the embryo immediately before hatching. Corresponds to character 12 in Vinther, Parry, Briggs, & Van Roy (2017).

Amathia: Reed & Cloney (1982).
Clupeafumosus socialis: Described by Topper et al. (2013R).
Conocardium elongatum: Pojeta, Runnegar, Morris, & Newell (1972).
Dentalium: The shell does not form until the trochophore larval stage, which has been exquisitely described in Antalis (Wanninger & Haszprunar, 2001).
This shell field is initially disc-like, subsequently expanding to fuse ventrally and produce the cylindrical protoconch. The prototroch is clearly delineated fro the telotroch in post-metamorphic juveniles (Wanninger & Haszprunar, 2001).
Gasconsia: The earliest shell is not described by Hanken & Harper (1985) or Watkins (2002).
Haliotis: Auzoux-Bordenave et al. (2010).
Loxosomella: Absent, with no possible equivalent (Nielsen, 1966).
Mytilus: Kniprath (1980).
Namacalathus: Inapplicable insofar as reproduction occurs by budding; there is no evidence for a free-living larval stage. Nevertheless, the presence of a sexual reproductive phase in addition to asexual reproduction cannot be discounted.
Neopilina: Not coiled, as stated in Lemche & Wingstrand (1959), but bulbous (Lindberg, 1985,Wingstrand (1985)).
Novocrania: Shell not secreted until after metamorphosis (Popov et al. 2010). Freeman & Lundelius (1999) report a Craniops-like larval shell in fossil “Crania”, but observe that Quaternary [Novo]crania no longer exhibit a larval shell.
Paramicrocornus: “The initial part of the conch appears to be a simple apex without clearly delineated protoconch” (Z.-L. Zhang, Skovsted, & Zhang, 2018), though it is not clear from illustrated figures whether an embryonic shell contiguous with the adult shell was present.
Pelagiella: Present (Nützel, Lehnert, & Frýda, 2006).
Pojetaia runnegari: Prodissoconch I probably demarcated by first growth line (B. Runnegar & Bentley, 1983).

Tonicella: On hatching, the polyplacophoran larva lacks a shell field.

Shell fields develop during the trochophore larva stage. The larva of the chiton Mopalia has two distinct shell fields: that anterior to the prototroch will develop into the first shell plate; the one posterior to the prototroch becomes the subsequent plates (Wanninger & Haszprunar, 2002a).

This disc-shaped posterior plate, whose position corresponds to the conchiferan shell field, bears a polygonal ornament and is subdivided by a series of grooves that prefigure the adult shell plates (Wanninger & Haszprunar, 2002a).
Yilingia spiciformis: Chen, Zhou, Yuan, & Xiao (2019).

[2] Morphology

Character adds 8 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[3]Neopilina[1]Conocardium elongatum[3]Haliotis[1]Pojetaia runnegari[1]Mytilus[-]Dentalium[5]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[3]Haplophrentis carinatus[1]Cupitheca holocyclata[3]Bactrotheca[1]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[-]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[-]Orthis[-]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[1]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[2]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[2]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 2: Brephic shell: Morphology

1: Flat, disc-like (cf. Micrina)2: Three prominent lobes forming a Y (cf. Paterimitra)3: Spherical5: High, conical?: Ambiguous-: InapplicableTransformational character.

The brephic shell is the shell possessed by the young organism (see Ushatinskaya & Korovnikov 2016R and Popov et al. 2010 for discussion of terminology).

Micrina resembles linguliforms (Holmer et al. 2011): in both, the brephic mitral shell has one pair of setal sacs enclosed by lateral lobes, whereas the brephic ventral shell has two lateral setal tubes.

Paterimitra and Salanygolina have “identical” ventral brephic shells (Holmer et al 2011), resembling the shape of a ship’s propeller.

Haplophrentis is coded following typical hyoliths, which have a spherical brephic shell; Pedunculotheca’s, in contrast, is seemingly cap-shaped.

Askepasma toddense: Renoid – see fig. 4B3 in Topper et al. 2013T.
Clupeafumosus socialis: The flat larval shell of Clupeafumosus resembles that of Micrina in outline (Topper et al. 2013R; cf. Holmer et al. 2011).
Conocardium elongatum: Pojeta et al. (1972).
Coolinia pecten: See fig. 3 in Bassett et al. 2017.
Craniops: The embryonic shell is more or less circular in outline – see Freeman & Lundelius, 1999, fig. 6A.
Cupitheca holocyclata: The impression of the larval shell on the operculum is flat and disc-like (Skovsted et al., 2016). The fusiform ‘protoconch’ (H.-J. Sun, Malinky, Zhu, & Huang, 2018) likely represents a larval (rather than brephic) shell.
Haliotis: Subspherical (Auzoux-Bordenave et al., 2010).
Lingula: See fig. 159 in Williams et al. 1997.
Lingulellotreta malongensis: Disc-like (G.-X. Li & Holmer, 2004).
Mickwitzia muralensis: Trifoliate appearance results from prominent attachment rudiment and bunching of setal sacs (Balthasar 2009T).
Micromitra: Subtriangular – essentially round.
Mytilus: Flat (though split into two via non-mineralized ligament) (Kniprath, 1980).
Neopilina: Appears cap-shaped and relatively flat in Lindberg (1985), but more bulbous in Wingstrand (1985).
Pelagodiscus atlanticus: See e.g. fig 169 in Williams et al. (1997).
Pojetaia runnegari: Flat and disc-like – though bivalved (B. Runnegar & Bentley, 1983).
Tonicella: Disc-like, subdivided by transverse grooves (Wanninger & Haszprunar, 2002a).

[3] Embryonic shell extended in larvae

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[2]Conocardium elongatum[2]Haliotis[2]Pojetaia runnegari[2]Mytilus[-]Dentalium[2]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[2]Haplophrentis carinatus[2]Cupitheca holocyclata[2]Bactrotheca[12]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[2]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[-]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[-]Orthis[-]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[2]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[2]Mickwitzia muralensis[?]Mummpikia nuda[2]Botsfordia[2]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[2]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[2]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 3: Brephic shell: Embryonic shell extended in larvae

1: Not extended; embryonic shell contiguous with adult shell2: Extended into larval shell, separated from adult shell by prominent nick-: InapplicableTransformational character.

Many taxa add to their embryonic shell (the protegulum possessed by the embryo upon hatching) during the larval phase of their life cycle. The shell that exists at metamorphosis, marked by a halo or nick point, is variously termed the “first formed shell”, “metamorphic shell” or “larval shell” (Bassett & Popov 2017).

Bactrotheca: There is a small ridge and a change in surface ornament at the end of the larval shell (Dzik, 1980).
Clupeafumosus socialis: Described by Topper et al. (2013R).
Craniops: Prominent nick; see Freeman & Lundelius 1999, fig. 6A.
Cupitheca holocyclata: Prominent nick (Skovsted et al., 2016).
Eoobolus: Nick point indicated by arrows in fig. 1 of Balthasar (2009T).
Haliotis: Auzoux-Bordenave et al. (2010).
Lingulellotreta malongensis: No prominent nick point (Holmer, Popov, Koneva, & Rong, 1997; G.-X. Li & Holmer, 2004).
Mytilus: Prominent nick (Martel, 2000).
Neopilina: “delimited from the surrounding adult shell by the innermost (first) concentric ridges of the periostracum” (Wingstrand, 1985).
Paramicrocornus: Not clearly delineated (Z.-L. Zhang et al., 2018), so either inapplicable or undifferentiated.
Pedunculotheca diania: The flattened region at the umbo of the ventral valve in smaller specimens conceivably represents an embryonic shell, though it may alternatively represent a cicatrix or colleplax-like structure.
Pelagiella: Nützel et al. (2006).
Tonicella: Wanninger & Haszprunar (2002a).

[4] Surface ornament

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[-]Dentalium[1]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[4]Micromitra[3]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[-]Tomteluva perturbata[1]Salanygolina[-]Terebratulina[-]Orthis[-]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[1]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[2]Botsfordia[2]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[2]Clupeafumosus socialis[1]Micrina[3]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 4: Brephic shell: Surface ornament

1: Smooth2: Rounded pits3: Polygonal impressions4: Pustulose?: Ambiguous-: InapplicableTransformational character.

Pitting of the larval shell characterises acrotretids and their relatives. Pustules occur on Paterinidae. See Character 3 in Williams et al. (2000) tables 5–6.

Askepasma toddense: Indented with hexagonal pits (Williams et al. 1998T, appendix 2).
Clupeafumosus socialis: “Larval shells on both valves […] are covered by fine, shallow pits” – Topper et al. 2013R.
Cupitheca holocyclata: Perfectly smooth (Skovsted et al., 2016).
Eoobolus: Pitted (Williams et al. 2000, table 8).
Haliotis: Auzoux-Bordenave et al. (2010).
Lingula, Pelagodiscus atlanticus: Smooth, following family-level codings of Williams et al. 2000, table 6.
Lingulellotreta malongensis: Smooth (Holmer et al., 1997; G.-X. Li & Holmer, 2004).
Micrina: Smooth (Holmer et al. 2011F).
Micromitra: Pustolose in Paterinidae (Williams et al. 2000, table 6).
Mytilus: Kniprath (1980).
Paterimitra: Polygonal texture present (Holmer et al. 2011F), as in the adult shell.
Pelagiella: Nützel et al. (2006).
Pojetaia runnegari: (???).
Salanygolina: Smooth (Holmer et al. 2009T).
Siphonobolus priscus: “Smooth brephic shell” – Popov et al. 2009.

[5] Larval attachment structure

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[-]Dentalium[1]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[-]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[-]Orthis[-]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[1]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[2]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[2]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[1]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 5: Brephic shell: Larval attachment structure

1: Without evidence of pedicle2: With evidence of pedicle?: Ambiguous-: InapplicableTransformational character.

Embryonic shells of Micrina and certain linguliforms exhibit a transversely folded posterior extension that speaks of the original presence of a pedicle in the embryo.

This is independent of the presence of an adult pedicle, which may arise after metamorphosis.

Clupeafumosus socialis: The larval shell embraces the pedicle foramen, suggesting a larval attachment. See fig. 4 of Topper et al. (2013R).
Cupitheca holocyclata: A pedicle is not inferred by Skovsted et al. (2016), and is difficult to reconcile with the apical morphology documented by H.-J. Sun et al. (2018).
Eoobolus: Lobe related to the attachment rudiment (Balthasar 2009T, fig. 2).
Haliotis: Auzoux-Bordenave et al. (2010).
Lingulellotreta malongensis: The pedicle foramen intersects the brephic shell (Holmer et al., 1997; G.-X. Li & Holmer, 2004), suggesting larval attachment.
Mickwitzia muralensis: Note the posterior lobe related to the attachment rudiment in fig. 2 of Balthasar 2009T.
Siphonobolus priscus: Interpreted as having planktotrophic (and thus non-attached) larvae (Popov et al. 2009).

[6] Setulose

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[0]Eccentrotheca[?]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 6: Brephic shell: Setulose

0: No evidence of setae in embryonic shell1: Setae presentNeomorphic character.

The protegulum of Micrina is penetrated with canals that were originally associated with setae, a character that it has in common with linguliforms (Holmer et al. 2011).

Botsfordia: “One specimen shows fine capillae running laterally from the posterior tubercles on the dorsal valve (Pl. 3, fig. 5b). This is possibly the imprints of setae.” – Ushatinskaya & Korovnikov 2016R.
Clupeafumosus socialis: Setal bundles interpreted as present in acrotretids by Ushatinskaya (2016P).
Cupitheca holocyclata: Absent (Skovsted et al., 2016).
Lingulellotreta malongensis: Possible suggestion of setal sacs present on brephic shell (Holmer et al., 1997; G.-X. Li & Holmer, 2004), but outline inadequately preserved to code with confidence; treated as ambiguous.
Mickwitzia muralensis: Four setal sacs.

[7] Setal sacs

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[0]Eccentrotheca[?]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 7: Brephic shell: Setal sacs

0: Absent1: PresentNeomorphic character.

Setal sacs are recognizable as raised lumps on the juvenile shell (see Bassett and Popov 2017).

Micrina and linguliforms have setal sacs on their mitral/dorsal embryonic shell, whereas these are absent in Paterimitra (Holmer et al 2011).

Bactrotheca: Following Bactrotheca (Dzik, 1980).
Botsfordia: A single pair of low tubercles are (Ushatinskaya & Korovnikov 2016R state “may be”) located in the middle region of the dorsal and the ventral brephic valve; these are interpreted as a single pair of setal sacs, with the identity of the (dorsally unpaired) tubercles uncertain.
Clupeafumosus socialis: Setal bundles interpreted as present in acrotretids by Ushatinskaya (2016P).
Lingula: Lingulids’ larval setae are not arranged in bundles (Carlson 1995).
Lingulellotreta malongensis: Possible suggestion of setal sacs present on brephic shell (Holmer et al., 1997; G.-X. Li & Holmer, 2004), but outline inadequately preserved to code with confidence; treated as ambiguous.
Novocrania, Pelagodiscus atlanticus: Three pairs (Carlson 1995).

[8] Number

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[-]Eccentrotheca[?]Orthrozanclus[2]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[2]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[3]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[3]Pelagodiscus atlanticus[2]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[2]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[2]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[2]Clupeafumosus socialis[1]Micrina[-]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 8: Brephic shell: Setal sacs: Number

1: One pair2: Two pairs3: Three pairs?: Ambiguous-: InapplicableTransformational character.

Two pairs on e.g. Coolina; one on e.g. Micrina.

Botsfordia: “larval shell with one to three apical tubercles in ventral valve and two in dorsal valve” (Williams et al. 2000) – if these correspond to setal sacs, then we interpret this as equivalent to one pair.

In B. minuta, the ventral valve bears a single medial tubercle (which in figured material seems to have two bilaterally symmetrical fields), whereas the dorsal valve bears two apical tubercles (G.-X. Li & Holmer, 2004) – supporting the interpretation of a single pair of setal sacs.
Clupeafumosus socialis: Two pairs identified in acrotretids by Ushatinskaya (2016P).
Mickwitzia muralensis: See fig. 2 in Balthasar 2009T.
Novocrania, Pelagodiscus atlanticus: Three pairs (Carlson 1995).
Siphonobolus priscus: Two pairs of setal sacs (Popov et al. 2009).

4.2 Larval setae: Paired bundles [9]

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[?]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 9: Larval setae: Paired bundles

0: Absent1: Present?: AmbiguousNeomorphic character.

Annelid chaetae are equivalent to the bundled setae expressed in certain brachiopod larvae. See character 12 in Vinther, Van Roy, & Briggs (2008).

Amathia: (Reed & Cloney, 1982).
Chaetoderma: In transverse rows, with setae seeming to arise from papillae (???).
Flustra: Absent (Zimmer & Woollacott, 2013).
Haliotis: The spines that adorn the cephalic tentacles (Auzoux-Bordenave et al., 2010) are conceivably homologous with setae.
Mopalia: Not bundled (???).
Serpula: Lüter (2000) contends that annelid larvae lack setae, but chaetae are present on day 16 of the larval development of Serpula (Keay, 2007).
Terebratulina: Williams et al. (1997).

4.3 Adult setae [10]

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[1]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[1]Pelagiella[1]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[1]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[?]Orthis[1]Alisina[1]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 10: Adult setae

0: Absent1: Present?: AmbiguousNeomorphic character.

Lüter (2000) demonstrates that the setae of larval and adult brachiopods exhibit fundamental structural differences and are conceivably not homologous structures. Larval setae are thus described separately.

Although preservation of setae in fossil brachiopods is exceptional, their presence can be inferred from shelly material (see Holmer, Skovsted, & Brock (2006)).

The girdle elements of aculiferan molluscs include chitinous material that is secreted by microvilli; following Vinther et al. (2017), these are coded as potential homologues of setae.

Acaenoplax hayae: The spines that adorn the ridges (???) are interpreted as equivalent to polyplacophoran girdle elements.

Acanthotretella spinosa: Note that the setae do not obviously emerge from tubes, leading Holmer and Caron to question their homology with the setae of other taxa (Heliomedusa, Mickwitzia).

Both valves of Acanthotretella were covered by long spine-like and shell penetrating setae. The setae of A. decaius are usually preserved along anterior and anterolateral margins (Hu et al. 2010).
Amathia: The teeth of the Bryozoan gizzard have been homologized with annelid setae (Gordon, 1975).
Clupeafumosus socialis: Setal bundles interpreted as present in acrotretids by Ushatinskaya (2016P).
Flustra: A gizzard is not present in all bryozoans, and has not been reported in Flustra.
Kulindroplax perissokomos, Phthipodochiton thraivensis, Glaphurochiton carbonarius: The girdle elements of aculiferan molluscs include chitinous material that is secreted by microvilli; following Vinther et al. (2017), these are coded as potential homologues of setae.
Haplophrentis carinatus: Not observed (Moysiuk, Smith, & Caron, 2017), despite suitable preservation.
Leptochiton: Kaas (1981).
Lingulellotreta malongensis: “Setae appear short, delicate, and are closely fringed with the entire
mantle margin, hardly extending beyond the edge of shell” – Zhang, Shu, Han, & Liu (2005).
Mopalia: Leise (1986).
Novocrania: “Adult craniids are without setae (a feature shared with the thecideides, the
shells of which are also cemented).” – Williams et al. 2007.

Orthrozanclus: The sclerites of Orthrozanclus are interpreted as being homologous to those of Halkieria.

Orthrozanclus occurs in preservational regimes that preserve sclerites in annelids and Wiwaxia, so additional seta-like sclerites – whose presence cannot be evaluated in Halkieria – are taken to be genuinely absent.
Pelagiella: Reported by Thomas & Vinther (2012).

Polysacos vickersianum: The spinose sclerites of multiplacophorans are generally considered to represent modified shell plates rather than girdle elements (???, Conway Morris (2006)).

(???) argues that the spines of Polysacos are homologous with polyplacophoran girdle elements.

However, aesthete canals form by the inclusion of the secretory mantle within the growing valve (Baxter, Jones, & Sturrock, 1987), which points to a fundamentally non-seta-like growth mechanism; rather than secretion by basal microvilli, multiplacophoran spines evidently grow by basal accretion without periodic replacement.

As such, the existence of girdle elements homologous to setae is not demonstrated by available fossil material.

Siphogonuchites multa: The nature of the spicules that constitute the Siphogonuchites shell is uncertain. We treat them here as homologous to chiton girdle elements, following Conway Morris & Chapman (1996), Bengtson (1992) and Vinther et al. (2017).

An equivalence to halkieriid sclerites is not apparent: sclerites must have been added to the edge of the Siphogonuchitid shell during growth, requiring an increase in the number of sclerite ‘rows’; and they do not follow a quincuncial arrangement in a straightforward manner.

The internal ornament of parallel lines (Bengtson, 1992; Conway Morris & Chapman, 1996) recalls the longitudinal chambers within microvillar-secreted setae, but occur on the inner surface of phosphatized chambers, so probably have a different origin.
Siphonobolus priscus: Phosphatised setae emerge from hollow spines (Popov et al. 2009).
Sipunculus: The absence of chitin or microvillar lineations in sipunculan hooks argues against their interpretation as setae, but they are coded as conceivable homologues, with these characteristics treated separately.

Tonicella: The girdle elements of certain polyplacophorans are chitinous and secreted by microvilli (Fischer, Maile, & Renner, 1980; Leise, 1988; Leise & Cloney, 1982); it is therefore likely that they are homologous with the setae of other lophotrochozoans.

They are not homologous with the shell; they exhibit a distinct mode of secretion and have a different organic scaffold (Treves, Traub, Weiner, & Addadi, 2003, Ehrlich (2010)).
Wiwaxia corrugata: Sclerites likely correspond with lophotrochozoan setae (Butterfield, 1990; Smith, 2014; Zhang, Smith, & Shu, 2015).

[11] Secretion

Character adds 3 to tree score (2 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[2]Sipunculus[1]Wirenia[2]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[1]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 11: Adult setae: Secretion

1: By basal microvilli2: Epicuticular-: InapplicableTransformational character.

The majority of lophotrochozoan sclerites bear a characteristic striated texture that denotes their secretion by basal microvilli (Butterfield, 1990). The seta-like hooks of sipunculans lack this texture, suggesting that they may not be homologous with other setae.

Canadia spinosa: Butterfield (1990).
Chaetoderma: Glands, associated with papillae, “presumed to secrete the calcareous spicules” (???).
Leptochiton: (???).
Mopalia: Leise (1986).
Sipunculus: No evidence of microvillar secretion (e.g. Schulze, Cutler, & Giribet, 2005).
Wirenia: Secreted from base in Epimenia (Okusu, 2002), presumably by microvilli.

[12] Microvillar diameter

Character adds 2 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[-]Sipunculus[?]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[1]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 12: Adult setae: Secretion: Microvillar diameter

1: Uniform2: Decreasing towards seta margin-: InapplicableTransformational character.

The diameter of secretory microvilli may vary across the diameter of a seta (Smith, 2014).

Amathia: No trend in microvillar size (Gordon, 1975).
Capitella: Schweigkofler, Bartolomaeus, & Salvini-Plawen (1998).
Leptochiton: (???).
Lingula: Widest in centre (Lüter, 2000).
Mopalia: Leise (1986).
Pelagodiscus atlanticus: Slight decrease towards margin in Discinisca (Lüter, 2003).
Serpula: Following Scolelepis (Hausen, 2005); Diasoma (Orrhage, 1971).
Terebratulina: Decreases towards margin in Terebratalia larvae (Gustus & Cloney, 1972).
Tonicella: Uniform (Fischer et al., 1980; Leise & Cloney, 1982).

[13] Microvillar canal aspect

Character adds 2 to tree score (1 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[-]Sipunculus[?]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[2]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 13: Adult setae: Secretion: Microvillar canal aspect

1: Round2: Polygonal; close-packed?: Ambiguous-: InapplicableTransformational character.

Lüter (2000) distinguishes between the polygonal outline of microvillar canals in adult brachiopod setae and the oval outline of larval setae.

Amathia: Polygonal (Gordon, 1975).
Capitella: Predominantly round, though becoming close packed in places (Schweigkofler et al., 1998).
Leptochiton: (???).
Lingula: Polygonal (Lüter, 2000).
Mopalia: Leise (1986).
Pelagodiscus atlanticus: Reported as hexagonal in Discina (Lüter, 2000).
Serpula: Annelid setae are prominently round, with gaps between each microvillar chamber (Orrhage, 1971).
Terebratulina: Polygonal in Calloria (Lüter, 2000).
Tonicella: Round (Fischer et al., 1980).
Wiwaxia corrugata: The loose spacing of pyrite infills of microvillar canals in Wiwaxia sclerites (Smith, 2014) argues against a close-packed arrangement.

[14] Organic constituent

Character adds 3 to tree score (2 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[1]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 14: Adult setae: Composition: Organic constituent

1: Chitin2: Horny protein-: InapplicableTransformational character.

The majority of lophotrochozoan sclerites are chitinous, occasionally hosting secondary biominerals.

Canadia spinosa, Wiwaxia corrugata: Presumed chitin due to preservational character.
Leptochiton: “The spicules and scales […] are composed of aragonite and a highly glycosylated proteinaceous organic matrix” (???; Treves et al., 2003).
Mopalia: Chitinous (Leise, 1986).
Sipunculus: Enzymatic test for chitin proved negative (Rice, 1993).

[15] Enamel

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[0]Capitella[?]Canadia spinosa[?]Sipunculus[0]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 15: Adult setae: Composition: Enamel

0: Absent1: Present?: AmbiguousNeomorphic character.

Certain setae are encapsulated in a 20 nm wide electron dense layer, termed “enamel” by Gustus & Cloney (1973). Enamel may be absent in larval setae (Lüter, 2003); this character refers to the condition in adult setae.

Amathia: Not evident (Gordon, 1975).
Capitella: Not evident (Schweigkofler et al., 1998).
Mopalia: Not visible in TEM sections (Leise, 1986).
Pelagodiscus atlanticus: Enamel layer apparent in Discina (Williams et al., 1997, fig. 47.1).
Serpula: Present in Nereis (Gustus & Cloney, 1973).
Terebratulina: Present in the terebratulid Calloria (and the Rhynchonellid Notosaria) (Lüter, 2000).
Tonicella: Not evident (Fischer et al., 1980; Leise, 1988).

[16] Mineralized core

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[0]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[1]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 16: Adult setae: Composition: Mineralized core

0: Absent1: PresentNeomorphic character.

Per character 4 in Vinther et al. (2017), the sclerites and spicules of many aculiferans have a calcareous core.

Acaenoplax hayae: The spines are ‘probably aragonitic’ (???).
Mopalia: Spicules in medial bundles (Leise, 1986) not considered equivalent.
Tonicella: “Small, simple hairs” (Leise, 1988).
Wirenia: Aragonitic (Okusu, 2002).

[17] Distribution

Character adds 7 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[3]Serpula[3]Capitella[3]Canadia spinosa[-]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[3]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[3]Pelagiella[1]Siphogonuchites multa[3]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[4]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[2]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[2]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[2]Alisina[2]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[1]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[1]Micrina[-]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 17: Adult setae: Distribution

1: Uniform 2: Only present at margins of shell3: In bundles, repeated on each metamere if serial repetition present4: In digestive tract?: Ambiguous-: InapplicableTransformational character.

Setae penetrate the valves of many brachiopods. In certain taxa, they are apparent only at the margins of the valves, in association with the commissure, being reduced or lost over the surface of the shell.

The ‘fascicles’ of Vinther et al. (2017) are a specific case of the ‘bundles’ described here.

Acaenoplax hayae: Strictly, in transverse rows (???), but in view of the serial repetition this state is deemed appropriate.
Calvapilosa kroegeri: No zonation evident (Vinther et al., 2017).
Chaetoderma: Repetition only evident in larvae (???).
Eccentrotheca: Skovsted et al (2011) assumed the setae may have been present along the margin of the adapical opening, but there is no fossil evidence.
Glaphurochiton carbonarius: Girdle at margin of organism (Hoare & Mapes, 1986).
Heliomedusa orienta: Throughout the shell – see Williams et al. 2007 – causing the pustulose appearance remarked upon by Chen et al. 2007.
Kulindroplax perissokomos: Sutton, Briggs, Siveter, Siveter, & Sigwart (2012).
Leptochiton: (???).
Lingulellotreta malongensis: At margin of shell (Zhang et al., 2005).
Mopalia: No obvious pattern or arrangement evident (Leise, 1986).
Phthipodochiton thraivensis: No pattern apparent (Sutton & Sigwart, 2012).
Siphogonuchites multa: No evidence of metamerism is evident (Bengtson, 1992).
Tonicella: Uniformly distributed around girdle (though not within shell) with no serial repetition (Leise, 1988; Vinther & Nielsen, 2005).
Wirenia: In Epimenia (Okusu, 2002).

[18] Present on anteriormost segment

Character adds 2 to tree score (1 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[1]Canadia spinosa[-]Sipunculus[?]Wirenia[?]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[?]Amathia[-]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 18: Adult setae: Distribution: Present on anteriormost segment

1: Chaetae present on prostomium2: Prostomium and peristomium lack chaetae?: Ambiguous-: InapplicableTransformational character.

This character attempts to reflect character 115 in L. A. Parry, Edgecombe, Eibye-Jacobsen, & Vinther (2016), as modified by (???). This character seeks to capture the fact that both Canadia and Phragmochaeta are interpreted as bearing chaetal bundles on their anterior segments (???). Wiwaxia does too.

I treat the character as transformational, coding it as inapplicable where trunk chaetae or parapodia are absent, as it is not possible to independently verify the ancestral state of this character.

[19] Internal constitution

Character adds 4 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[2]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[2]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[1]Pelagiella[2]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[2]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 19: Adult setae: Internal constitution

1: Solid, blade-like2: Basal invagination-: InapplicableTransformational character.

Sipunculan “setae” are basally invaginated, suggesting that they may not be homologous with annelid chaetae. Certain aculiferans also exhibit basally hollow sclerites (???, character 6).

Acaenoplax hayae: No indication of basal invagination (???).
Amathia: Cytoplasmic intrusion into a central cavity (Gordon, 1975).
Calvapilosa kroegeri: Hollow conical sclerites (Vinther et al., 2017).
Capitella: Schweigkofler et al. (1998).
Chaetoderma: (???).
Kulindroplax perissokomos: The sclerites are described as blade-like (Sutton et al., 2012); their concavo-convex shape presumably relates to the surface of the blade, rather than being intended to imply a deep basal cavity.
Mopalia: Leise (1986).
Pelagiella: Seemingly without invagination.
Phthipodochiton thraivensis: Inadequately preserved to determine (Sutton & Sigwart, 2012).
Siphogonuchites multa: Profoundly hollow: deep basal invagination (Bengtson, 1992; Conway Morris & Chapman, 1996).
Wirenia: Sealed base, despite internal chamber, in Epimenia (Okusu, 2002).

[20] Distinct shaft

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[0]Canadia spinosa[0]Sipunculus[1]Wirenia[0]Chaetoderma[0]Tonicella[1]Mopalia[?]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[1]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 20: Adult setae: Distinct shaft

0: Absent1: Present?: AmbiguousNeomorphic character.

The setae of certain taxa (e.g. Wiwaxia, Mopalia) have a differentiated shaft that inserts into the body wall.

Canadia spinosa: Butterfield (1990).
Capitella: Shaft and hood (Schweigkofler et al., 1998).
Chaetoderma: (???).
Glaphurochiton carbonarius: Seemingly absent (Hoare & Mapes, 1986).
Pelagiella: No shaft evident (Thomas & Vinther, 2012).
Siphogonuchites multa: Absent, in contradistinction to Halkieria (Conway Morris & Chapman, 1996).
Wirenia: Somewhat distinct in Epimenia (Okusu, 2002).
Wiwaxia corrugata: Smith (2014).

[21] Hooks

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[0]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 21: Adult setae: Hooks

0: Absent1: PresentNeomorphic character.

Hooked chaetae arise through the reorientation of the chaetoblast during secretion (Hausen, 2005). (???) and L. A. Parry et al. (2016) distinguish falcate (sickle-shaped) hooks (characters 121 and 98 respectively), dentate hooks (characters 122 and 92) and uncini (characters 123 and 94) as fundamentally different types of chaetae. A dentate hook, however, can be seen as a falcate hook with additional adrostral teeth or processes (???). We therefore code simply for the presence of apical curvature in any chaetae, with a view that a gain of a ‘hook’ represents an evolutionary novelty that might then be expressed in various locations and complemented by the addition of subsidiary dentition (i.e. adrostral hooks).

For terminology, see (???) and (???).

Capitella: Present in Notomastus (Capitellidae) (Hausen, 2005, fig. 4).
Serpula: Following coding of character 80 in (???); uncini scored as present by L. A. Parry et al. (2016).

[22] Capitium

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[0]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 22: Adult setae: Capitium

0: Absent1: PresentNeomorphic character.

Character 81 in (???). The capitium is a region on the convex surface of the rostrum (if present) that contains containing multiple teeth, each secreted by an individual microvillus (???, ???; Hausen, 2005), and in a consistent orientation with the margin of the chaeta.

Capitella: Present in Notomastus (Capitellidae) (Hausen, 2005, fig. 4).

[23] Projecting knobs

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[1]Capitella[1]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 23: Adult setae: Projecting knobs

0: Absent1: Individual peripheral projections present?: AmbiguousNeomorphic character.

Terebratulids and discinids instead exhibit knob-like individual spines. These are distinct from the rings of spines that fringe lingulid setae.

Calvapilosa kroegeri: Vinther et al. (2017).
Canadia spinosa: Marginal knobs and serrations (Butterfield, 1990).
Capitella: Projections around hood (Schweigkofler et al., 1998).
Chaetoderma: (???).
Lingula: Lüter (2000).
Pelagodiscus atlanticus: Discinisca sports individual peripheral spines (Lüter, 2003; Williams et al., 1997).
Note that the “embryonic” setae of Discinids correspond to the “larval setae” of other brachiopods, and the “larval setae” of juvenile discinids correspond to adult setae (Lüter, 2003).
Serpula: Surface smooth (Y. Sun & Qiu, 2012).
Siphogonuchites multa: Conway Morris & Chapman (1996).
Terebratulina: Individual peripheral spines (in Calloria; Lüter, 2000).
Wirenia: In Epimenia (Okusu, 2002).

[24] Circling coronae

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 24: Adult setae: Circling coronae

0: Absent1: PresentNeomorphic character.

Lingulid setae bear crown-like rings of fine spines delimiting vertical sections, recalling the nodes of Equisetum stems. These arise by the addition of an additional circlet of microvilli (see Lüter, 2000, fig. 1e).

Capitella: Schweigkofler et al. (1998).
Chaetoderma: (???).
Wirenia: Epimenia (Okusu, 2002).

4.4 Body organization

[25] Serial repetition

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[1]Sipunculus[0]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 25: Body organization: Serial repetition

0: Absent1: Present?: AmbiguousNeomorphic character.

Serial repetition in adult, whether expressed in valves, soft tissues or exoskeletal elements. See character 13 in Rouse (1999); 19 in Vinther et al. (2008); 38 in Haszprunar (1996); 40–41 in Sutton & Sigwart (2012); Wanninger (2009).

Acaenoplax hayae: Serial repetition of lobes and spine rows (???).
Bactrotheca: The soft anatomy of this taxon is unknown, so it is impossible to rule out the presence of serial repetition.
Calvapilosa kroegeri: Conceivably evident in soft tissue that is not preserved, e.g. gills.
Chaetoderma: Present in larval stages only (???).
Dailyatia: Unknown whether sclerites are serially repeated, or whether metameres were present in underlying soft anatomy.
Halkieria evangelista: Elements of the Halkieria scleritome adhere to a quincunx arrangement, with different spacing of elements in each zone; there is no evidence of a metameric arrangement.
Namacalathus, Wirenia: Not evident.
Neopilina: Serially repeated gills.
Odontogriphus omalus: Repetition of ctenidia (???).
Orthrozanclus: Coded following Halkieria.
Pelagiella: Not apparent in distribution of setae (Thomas & Vinther, 2012).
Yilingia spiciformis: Present (Chen et al., 2019).

[26] Annulae

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[0]Canadia spinosa[1]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 26: Body organization: Annulae

0: Absent1: Prominent annulation of epidermis?: AmbiguousNeomorphic character.

The trunk of Sipunculus and many annelid worms bear annulations.

[27] Subdivided head

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[0]Canadia spinosa[1]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 27: Body organization: Subdivided head

0: Undivided1: Comprising prostomium and peristomium?: AmbiguousNeomorphic character.

The annelid head comprises a differentiated prostomium and peristomium. See character 119 in Parry & Caron (2019).

Canadia spinosa, Serpula: Following Parry & Caron (2019).
Capitella: Fused prostomium and peristomium, following Parry & Caron (2019).
Halkieria evangelista: Unknown (contra Parry & Caron, 2019); no soft tissue preservation.
Odontogriphus omalus, Wiwaxia corrugata: No evidence of subdivision.
Sipunculus: The sipunculan proboscis corresponds to the annelid prostomium (???).

[28] Pedal groove

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[1]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 28: Body organization: Pedal groove

0: Absent1: PresentNeomorphic character.

Certain aculiferans have secondarily lost a foot, but retain a pedal groove.

Acaenoplax hayae: (???).
Kulindroplax perissokomos: Sutton et al. (2012).
Phthipodochiton thraivensis: Seemingly present, though foot unequivocally absent (Sutton & Sigwart, 2012; Sutton et al., 2012).
Wirenia: Present and flanked by a distinct set of spicules in Epimenia (Okusu, 2002).

[30] Coelom

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 30: Body organization: Coelom

0: Absent: adults acoelomate1: Present: true coelomic cavities differentiatedNeomorphic character.

Bactrotheca: The soft anatomy of this taxon is unknown, so it is impossible to determine the presence of a coelom.
Flustra: “Adult ectoprocts differentiate true coelomic cavities” – Fuchs & Wanninger (2008).
Haplophrentis carinatus: Internal cavities indicated by differentiation of internal organs (see Moysiuk et al., 2017).
Loxosomella: “Adult entoprocts are acoelomate” – Fuchs & Wanninger (2008).
Phoronis: Temereva (2017).

[31] Number

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[-]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 31: Body organization: Coelomoducts: Number

1: Single2: Multiple?: Ambiguous-: InapplicableTransformational character.

Character 27 in Haszprunar (2000). Coelomoducts are excretory organs derived from the coelom, also in some cases serving as genital ducts (gonoducts); they replace (and may resemble) nephridia (Goodrich, 1945).

Flustra: Multiple ciliated ducts leading to a common gonopore (Goodrich, 1945).
Loxosomella: Two coelomoducts pass outwards, meet, and open by a common pore (Goodrich, 1945).
Phoronis: “large coelomic funnels serving as genital ducts” (Goodrich, 1945).

[32] Gills

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[1]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 32: Body organization: Gills

0: Absent1: PresentNeomorphic character.

Gills (or ctenidia) surround the molluscan foot.
Character 1.59–60, 2.09, 4.49 in von Salvini-Plawen & Steiner (1996); 10–11 in Haszprunar (2000); 45 in Sutton & Sigwart (2012).

Acaenoplax hayae: The posterior projections are interpreted as resembling the gill folds of solenogasatres (???).
Calvapilosa kroegeri: Not preserved, though presumably present.
Canadia spinosa: Parry & Caron (2019).

[33] Ctenidia

Character adds 4 to tree score (3 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[2]Canadia spinosa[-]Sipunculus[1]Wirenia[2]Chaetoderma[2]Tonicella[2]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[2]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[2]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 33: Body organization: Gills: Ctenidia

1: Gills without filaments or leaflets2: Ctenidia: possessing filaments or leaflets-: InapplicableTransformational character.

Acaenoplax hayae: The posterior projections are interpreted as resembling the gill folds of solenogasatres (???).
Canadia spinosa: Parry & Caron (2019).
Wirenia, Chaetoderma: “Gill folds” in solenogastres; true ctenidia in caudofoveates (???).
Kulindroplax perissokomos: “More closely comparable to the ctenidia of caudofoveates” (Sutton et al., 2012).
Odontogriphus omalus: Gills appear to have leaflets (???).
Wiwaxia corrugata: Unclear (Smith, 2014).

[34] Circulatory system

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[3]Serpula[4]Capitella[?]Canadia spinosa[1]Sipunculus[2]Wirenia[2]Chaetoderma[2]Tonicella[2]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[2]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 34: Body organization: Circulatory system

1: Epithelially lined2: Poorly defined, involving sinuses and lacunae3: Closed circulatory system4: Blood within mesenteria of coelomic cavities?: AmbiguousTransformational character.

After character 23 in Haszprunar (1996); 24 in Haszprunar (2000); 41 in Rouse (1999); 16 in A. H. Scheltema (1993); 16 in Vinther et al. (2008); 5 in Haszprunar & Wanninger (2008).

Flustra, Amathia: As Brachiopods, sipunculans and relatives (Ruppert & Carle, 1983).
Loxosomella, Tonicella, Dentalium: See Haszprunar & Wanninger (2008).
Sipunculus: Open circulatory system.

4.5 Pedicle

[35] Presence

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[1]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[?]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[?]Clupeafumosus socialis[1]Micrina[1]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 35: Pedicle: Presence

0: Absent1: PresentNeomorphic character.

The brachiopod pedicle is a fleshy protuberance that emerges from the posterior part of the body wall – as denoted in fossil taxa by its occurrence between the dorsal and ventral valves.

It is important to distinguish the pedicle from the “pedicle sheath”, a tubular extension of the umbo that grows by accretion from an isolated portion of the ventral mantle. For discussion see Holmer et al. 2018T and Bassett and Popov 2017.

Acanthotretella spinosa: The attachment structure of Acanthotretella originates at the margin of the dorsal and ventral valves; although it emerges from the umbo of the ventral valve, the presence of an internal pedicle tube betrays its identity as a pedicle, rather than a pedicle sheath.

The pedicle of Acanthotretella emerges from a short extension of the umbo of the ventral valve. This extension is contiguous with the valve and presumably grew by accretion; its position and continuity with the valve suggest its interpretation as a pedicle sheath that is superseded as an attachment structure. On the other hand, its continuity with the internal pedicle tube suggests that is may represent an independent organ.
Bactrotheca: The apex of Bactrotheca deleta is pointed (Novak, 1891).
Botsfordia: Pedicle foramen was not necessarily occupied by a pedicle (though it presumably was).
Clupeafumosus socialis: A pedicle was presumably present, but only the foramen is preserved.
Cotyledion tylodes: The stalk is conceivably homologous with the brachiopod pedicle, but this possibility is impossible to test.
Craniops: Attached apically by cementation.
Cupitheca holocyclata: Not possible to reconcile with decollation.
Flustra: Grows directly onto the substrate.
Heliomedusa orienta: “It seems unlikely that H. orienta possessed a pedicle that attached it to
the soft seafloor, like most other Chengjiang brachiopods.” …
“The putative pedicle illustrated by Chen et al. (2007: Figs 4, 6, 7) in fact is the mold of a three-dimensionally preserved visceral cavity” – Zhang et al. 2009.
Lingulosacculus: The absence of a pedicle is inferred from the absence of an internal pedicle tube, and the absence of a pedicle at the hinge.
Loxosomella: The stalk corresponds to the molluscan foot, rather than a pedicle.
Mickwitzia muralensis: An attachment structure is inferred based on the presence of an opening (Balthasar 2004); this is assumed to have been homologous with the brachiopod pedicle.
Micrina: The prominent foramen between artificially articulated valves is taken to imply the presence of a pedicle (Holmer, Skovsted, Brock, Valentine, & Paterson, 2008).
Micromitra: The presence of a pedicle is indicated by the propensity of Micromitra to attach to hard substrates, such as sponge spicules (Holmer & Caron, 2006).
Namacalathus: There is no obvious way to homologise the attachment structure with the ventral pedicle of brachiopods.
Nisusia sulcata: Has a pedicle, rather than a pedicle sheath as in Kutorgina (Holmer et al. 2018E; Holmer et al. 2018T).
Paterimitra: “Paterimitra is interpreted to have attached to hard substrates via a pedicle that emerged through the small posterior opening” – Skovsted et al. 2009.
Phoronis: The tube-bearing stalk of phoronids arises as an eversion of the metastomal sac, a markedly different origin from the brachiopod pedicle, which arises from a terminal attachment disc (Young, 2002); the structures are of dubious homology.
Siphonobolus priscus: Presumed present, based on ventral foramen with colleplax.
Sipunculus: Absent; there is no clear basis to homologise the larval attachment structure of certain sipunculans with a pedicle.

[36] Constitution

Character adds 4 to tree score (3 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[1]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[1]Orthis[2]Alisina[2]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 36: Pedicle: Constitution

1: Massive or uniform2: Densely stacked tabular discs-: InapplicableTransformational character.

The pedicle of certain chengjiang rhynchonelliforms comprises “densely stacked, three dimensionally preserved, tabular discs” (Holmer et al. 2018E).
This contrasts with the uniform (‘massive’) pedicles of living taxa.

Antigonambonites planus: Biomineralized (Holmer et al. 2018E).
Terebratulina: Extant rhynconellid pedicles are massive, consisting of a thick outer chitinous cuticle, a pedicle epithelium, and a core composed of collagen fibres and cartilage-like connective tissue (Holmer et al. 2018E).

[37] Biomineralization

Character adds 2 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[2]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[1]Micromitra[?]Askepasma toddense[?]Coolinia pecten[2]Antigonambonites planus[2]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[12]Alisina[1]Longtancunella chengjiangensis[?]Glyptoria[1]Nisusia sulcata[12]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[1]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 37: Pedicle: Biomineralization

1: Absent2: Present?: Ambiguous-: InapplicableTransformational character.

Kutorgina chengjiangensis: The tabular discs that make up the pedicle “clearly have a pronounced three-dimensional preservation and may have been partly mineralized.” –Holmer, Zhang, Topper, Popov, & Claybourn (2018b).
Micromitra: A pedicle has not been observed in biomineralized material (Williams, Popov, Holmer, & Cusack, 1998b), indicating an originally non-mineralized constitution.

[38] Bulb

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[0]Micromitra[?]Askepasma toddense[?]Coolinia pecten[0]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[0]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 38: Pedicle: Bulb

0: Absent1: Present?: AmbiguousNeomorphic character.

A bulb is an expanded region of the distal pedicle, often embedded into the sediment to improve anchorage.

Acanthotretella spinosa: Holmer and Caron (2006) interpret the presence of a bulb as tentative; we score it as ambiguous.

[39] Distal rootlets

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[1]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[0]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 39: Pedicle: Distal rootlets

0: Absent1: Present?: AmbiguousNeomorphic character.

Observed in Pedunculotheca and Bethia (Sutton et al 2005).

[40] Tapering

Character adds 5 to tree score (4 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[2]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[2]Antigonambonites planus[2]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[2]Alisina[2]Longtancunella chengjiangensis[?]Glyptoria[2]Nisusia sulcata[1]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 40: Pedicle: Tapering

1: Uniform thickness2: Tapering?: Ambiguous-: InapplicableTransformational character.

Holmer et al. (2018T) remark that the tapering aspect of the Nisusia pedicle recalls that of certain Chengjiang taxa (Alisina, Longtancunella) whilst distinguishing it from many other taxa (Eichwaldia, Bethia) in which the pedicle is a constant thickness.

Antigonambonites planus: Tapered pedicle sheath with holdfast.
Pedunculotheca diania: The pedicle thickness gradually typering from the apex of the shell to the holdfast.

[41] Coelomic region

Character adds 2 to tree score (1 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[2]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[2]Antigonambonites planus[2]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[1]Alisina[1]Longtancunella chengjiangensis[?]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[2]Mummpikia nuda[?]Botsfordia[?]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[2]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 41: Pedicle: Coelomic region

1: Absent2: Present?: Ambiguous-: InapplicableTransformational character.

Certain brachiopods, such as Acanthotretella, exhibit a coelomic cavity within the pedicle or pedicle sheath.

Treated as transformational as it is not clear that either state is necessarily ancestral.

Nisusia sulcata: A coleomic canal is inferred based on the ease with which the pedicle is deformed (Holmer et al. 2018E), but its presence is not known for certain so is coded ambiguous.

[42] Surface ornament

Character adds 4 to tree score (3 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[2]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[2]Antigonambonites planus[2]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[2]Alisina[2]Longtancunella chengjiangensis[?]Glyptoria[23]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[2]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[2]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 42: Pedicle: Surface ornament

1: Smooth2: Regular annulations?: Irregular wrinkles-: AmbiguousTransformational character.

Annulations are regular rings that surround the pedicle, and are distinguished from wrinkles, which are irregular in magnitude and spacing, and may branch or fail to entirely encircle the pedicle.

Acanthotretella spinosa: “The pedicle surface is ornamented with pronounced annulated rings, disposed at intervals of about 0.2 mm”.
Alisina: “It appears that the pedicle lacks a coelomic space and is distinctly annulated, with densely stacked tabular bodies” – Zhang et al. 2011A.
Antigonambonites planus: “The emerging pedicle has a consistent shape in all the available specimens and is strongly annulated and distally tapering” – Holmer et al. 2018E.
Kutorgina chengjiangensis: “Pronounced concentric annular discs disposed at intervals of 0.6–1.0 mm” – Zhang et al. (2007b).
Lingulellotreta malongensis: Regularly annotated (see fig. 14.9 in Hou et al. 2017).
Longtancunella chengjiangensis: “The preserved pedicle has condensed annulations” – Zhang et al. 2011T.
Nisusia sulcata: The “strong annulations” vary significantly in transverse thickness (Holmer et al. 2018E), so it is not clear whether these represent true annulations or wrinkles.
Yuganotheca elegans: Annulations present in median collar.

[43] Nerve impression

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[?]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[?]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[?]Heliomedusa orienta[0]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[0]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[0]Clupeafumosus socialis[?]Micrina[?]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 43: Pedicle: Nerve impression

0: Absent1: PresentNeomorphic character.

In certain taxa the impression of the pedicle nerve is evident in the shell. See character 28 in Williams et al. (1998T) appendix 1. Care must be taken not to code an impression as absent when the preservational quality is insufficient to safely infer a genuine absence. Treated as neomorphic as the presence of an innervation is considered a derived state.

Alisina: Not described by Williams et al. 2000.
Askepasma toddense, Micromitra, Glyptoria, Kutorgina chengjiangensis, Salanygolina: Following Williams et al. 1998T, appendix 2.
Botsfordia: Documented by Skovsted et al. 2017.
Clupeafumosus socialis: Coded as absent in Acrotretidae (Williams et al. 2000, table 6).
Lingula: Present in many lingulids (Williams et al. 2000), and coded as present in Lingulidae (Williams et al. 2000, table 6).
Lingulellotreta malongensis: Coded as present in Lingulellotretidae (Williams et al. 2000, table 6).
Mummpikia nuda: Balthasar (2008, p. 274) identifies a canal as a probable impression of a pedicle nerve.
Nisusia sulcata, Orthis: Not reported in Williams et al. 2000.
Pelagodiscus atlanticus: Coded as present in Discinidae (Williams et al. 2000, table 6).
Siphonobolus priscus: Coded as absent in Siphonotretidae (Williams et al. 2000, table 6).

4.6 Mantle cavity

[44] Presence

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 44: Mantle cavity: Presence

0: Absent1: PresentNeomorphic character.

Character 8 in Haszprunar (2000).

Loxosomella, Flustra: Absent: Mantle absent.

[45] Reduced

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 45: Mantle cavity: Reduced

0: Not reduced1: Reduction of mantle cavityNeomorphic character.

Reduced in aplacophorans; see A. H. Scheltema (1993).

[46] Open face

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[2]Wirenia[2]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[2]Acaenoplax hayae[2]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 46: Mantle cavity: Open face

1: Open face2: Blind sac-: InapplicableTransformational character.

The mantle cavity (pallial cavity) has an open face in polyplacophorans, but forms a blind sac in taxa such as gastropods. See (???).

Calvapilosa kroegeri: Extensive, open-faced mantle cavity surrounding foot (Vinther et al., 2017).
Phthipodochiton thraivensis: Indirectly implied (Sutton et al., 2012).

[47] Sac opening

Character adds 3 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[2]Wirenia[1]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[3]Haliotis[?]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 47: Mantle cavity: Sac opening

1: Posteriad2: Posteroventrally3: Ventrally-: InapplicableTransformational character.

Caudofoveate and solenogaster aplacophorans can be distinguished based on the direction that their mantle cavity opens (Sutton et al., 2012).

Wirenia, Chaetoderma, Kulindroplax perissokomos: Sutton et al. (2012).
Phthipodochiton thraivensis: Indirectly interpreted as posterioventral (Sutton et al., 2012).

[48] Pallial line

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[0]Haliotis[1]Pojetaia runnegari[1]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 48: Mantle cavity: Pallial line

0: Absent1: PresentNeomorphic character.

The pallial line is a mark on inner surface of a shell reflecting the attachment of the mantle.

Conocardium elongatum: Prominent in some rostroconchs (Runnegar, 1978).
Neopilina: Present, marking limit of nacreous layer (McLean, 1979).

4.7 Mantle canals

[49] Presence

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[?]Haplophrentis carinatus[0]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[?]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 49: Mantle canals: Presence

0: Absent1: Present?: AmbiguousNeomorphic character.

Whether impressed on a shell or expressed solely in soft tissue.

Cupitheca holocyclata: Not observed despite excellent preservation.
Paramicrocornus: Not impressed on valves, despite fine preservation of muscular attachment (Z.-L. Zhang et al., 2018).
Paterimitra: Not evident.
Ussunia: Not preserved along muscle scars (Nikitin & Popov, 1984), presumably owing to quality of preservation rather than genuine absence.

[50] Morphology

Character adds 7 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[?]Haplophrentis carinatus[-]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[4]Micromitra[1]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[4]Orthis[3]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[3]Botsfordia[3]Eoobolus[3]Lingulellotreta malongensis[2]Lingula[3]Siphonobolus priscus[3]Acanthotretella spinosa[3]Lingulosacculus[1]Clupeafumosus socialis[?]Micrina[-]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 50: Mantle canals: Morphology

1: Pinnate (=lemniscate)2: Bifurcate3: Baculate4: Saccate?: Ambiguous-: InapplicableTransformational character.

The morphology of dorsal and ventral canals is identical in all included taxa, so is assumed not to be independent – hence the use of a single character (contra Williams et al. 2000).

For a description of terms see Williams et al. (1997, 2000).

Pinnate = “rapidly branch into a number of subequal, radially disposed canals”
Bifurcate = “vascula lateralia in both valves divide immediately after leaving the body cavity”
Baculate = “extend forward without any major dichotomy or bifurcation” (Williams et al. 1997 p. 418)
Saccate = “pouchlike sinuses lying wholly posterior to the arcuate vascula media” (ibid., p412).

Acanthotretella spinosa: Following Table 6, for Siphonotretidae, in Williams et al. (2000).
Alisina, Nisusia sulcata: Following Table 15 in Williams et al. (2000).
Antigonambonites planus: Not reported in Treatise (Williams et al. 2000).
Askepasma toddense: Described as pinnate (at least in ventral valve) by Williams et al. (1998T, p. 250).
Botsfordia, Eoobolus: Following Williams et al. 1998T, appendix 2, and Williams et al (2000), table 8.
Clupeafumosus socialis: Following Table 8 (for Acrotreta) in Williams et al. (2000), and the general pinnate condition for acrotretoids stated in Williams et al. (1997), p. 420.
Coolinia pecten: Not reported in Williams et al. 2000.
Craniops: Not reported from fossil material.
Gasconsia: Williams et al. (2000, table 15) appear to use Palaeotrimerella (as drawn in Williams et al. 1997) as a model for Gasconsia, which pre-supposes a close relationship. We are not aware of any report of mantle canals from Gasconsia itself.
Glyptoria: Following appendix 2 (char. 21) in Williams et al. (1998T).
Heliomedusa orienta: Described as pinnate by Jin & Wang (1992).
Novocrania, Kutorgina chengjiangensis: Following table 15 in Williams et al. (2000) (for Neocrania).
Lingula, Lingulellotreta malongensis: Following table 6 in Williams et al. (2000).
Lingulosacculus: Baculate vascula media – Balthasar & Butterfield (2009E).
Longtancunella chengjiangensis: Reported by Zhang et al. (2007A, 2011T) though the interpretation is tentative.
Micromitra: Described as saccate by Williams et al. (1998T).
Mummpikia nuda: “Poorly resolved” – Balthasar 2008.
Orthis: Sacculate (sometimes digitate in dorsal valve) (Williams et al. 2000, p716).
Pelagodiscus atlanticus: Following table 6, for Discinidae, in Williams et al. (2000).
Salanygolina: Coded uncertain in appendix 2 in Williams et al. (1998T).
Siphonobolus priscus: Interpreted as baculate, following Havlicek 1982.
Terebratulina: “In modern terebratulides, the vascula media are subordinate to the lemniscate or pinnate vascula genitalia” – Williams et al. 1997.
Tomteluva perturbata: Preservation not adequate to evaluate (Streng 2016).

[51] vascula lateralia

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[?]Haplophrentis carinatus[0]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[?]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 51: Mantle canals: vascula lateralia

0: Absent1: Present?: AmbiguousNeomorphic character.

We treat the vascula lateralia as equivalent to the vascula genitalia of articulated brachiopods, allowing phylogenetic analysis to test their proposed homology.

Williams et al (1997) write: “The mantle canal system of most of the organophosphate-shelled species consists of a single pair of main trunks in the ventral mantle (vascula lateralia) and two pairs in the dorsal mantle, one pair (vascula lateralia) occupying a similar position to the single pair in the ventral mantle and a second pair projecting from the body cavity near the midline of the valve. This latter pair may be termed the vascula media, but whether they are strictly homologous with the vascula media of articulated brachiopods is a matter of opinion. It is also impossible to assert that the vascula lateralia are the homologues of the vascula myaria or genitalia of articulated species, although they are likely to be so as they arise in a comparable position.”

“In inarticulated brachiopods, two main mantle canals (vascula lateralia) emerge from the main body cavity through muscular valves and bifurcate distally to produce an increasingly dense array of blindly ending branches near the periphery of the mantle (fig. 71.1–71.2).”

Acanthotretella spinosa: Following table 8 (which records presence in Siphonotreta) in Williams et al. (2000).
Alisina, Kutorgina chengjiangensis, Nisusia sulcata: Following table 15 in Williams et al. (2000).
Askepasma toddense, Micromitra: “Laurie (1987) has shown that arcuate vascula media were present in the mantles of both valves as were pouchlike vascula genitalia, especially in the ventral valve” – Williams et al. 1997.
Botsfordia: Following Popov (1992).
Clupeafumosus socialis: Presence indicated in Table 8 (for Acrotreta) in Williams et al. (2000).
Gasconsia: Williams et al. (2000, table 15) appear to use Palaeotrimerella (as drawn in Williams et al. 1997) as a model for Gasconsia, which pre-supposes a close relationship. We are not aware of any report of mantle canals from Gasconsia itself.
Heliomedusa orienta: Present: Williams et al. (2000); Jin & Wang (1992).
Lingulellotreta malongensis: Present (Williams et al. 2000).

Longtancunella chengjiangensis: Presence is possible but requires interpretation that is not unambiguous:

“In the dorsal valve, there can be seen two baculate grooves that arise from the
anterior body wall at an antero-lateral position. These two grooves (Figs 4H, 5D) could be taken to represent the vascula lateralia” – Zhang et al 2007A.
Novocrania: Following table 15 in Williams et al. (2000) (for Neocrania), who write that “Holocene craniides have only a single pair of main trunks in both valves, corresponding to the vascula lateralia”. Williams et al. (2007) reiterate this position (p. 2875), at least for the ventral valve.
Terebratulina, Orthis: = vascula genitalia.
Pelagodiscus atlanticus: Following Lochkothele (Discinidae), Fig. 43.4a in Williams et al. (2000).
Siphonobolus priscus: Noted in Siphonobolus by Williams et al. (2000), with reference to Havlicek (1982).
Tomteluva perturbata: Preservation not adequate to evaluate (Streng 2016).
Yuganotheca elegans: Based on the figures and sketches in Zhang et al. 2014 (and supplementary material), the mantle canals are interpreted as lateral, with no clear vascula media present.

[52] vascula media

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[?]Haplophrentis carinatus[0]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[0]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[?]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 52: Mantle canals: vascula media

0: Absent1: Present (in dorsal valve)?: AmbiguousNeomorphic character.

Williams et al. (1997) note that in addition to the vascula lateralia, “Discinisca has two additional mantle canals emanating from the body cavity into the dorsal mantle (vascula media).”

These structures are only evident in the dorsal valve for the included taxa, so only a single character is necessary.

Acanthotretella spinosa: Following table 6 (for Siphonotretidae) in Williams et al. (2000).
Alisina, Kutorgina chengjiangensis, Nisusia sulcata: Following table 15 in Williams et al. (2000).
Askepasma toddense: Following table 6 (for Paterinidae) in Williams et al. (2000).
Botsfordia: Following Popov (1992, fig. 2).
Clupeafumosus socialis: Following Hadrotreta schematic in Williams et al. (2000).
Eoobolus: Fig. 5 in Balthasar 2009T.
Gasconsia: Williams et al. (2000, table 15) appear to use Palaeotrimerella (as drawn in Williams et al. 1997) as a model for Gasconsia, which pre-supposes a close relationship. We are not aware of any report of mantle canals from Gasconsia itself.
Glyptoria: Present and divergent (Williams et al. 2000).
Heliomedusa orienta: Present: Williams et al. (2000) p162, Jin & Wang (1992).
Lingula, Lingulellotreta malongensis: Following table 6 in Williams et al. (2000).
Longtancunella chengjiangensis: Reported by Zhang et al. (2007A) though the interpretation is tentative.
Micromitra: Reported by Williams et al. (1998T).
Novocrania: Williams et al. (2000) write “Holocene craniides have only a single pair of main trunks in both valves, corresponding to the vascula lateralia” – an observation reflected in their table 15 (for Neocrania).
But in contrast, Williams et al. 2007, p. 2875, identify the dorsal valve’s canals as a vascula media in living cranidds (though both are lateralia in Ordoviian craniides). This character is therefore coded as ambiguous.
Orthis: From idealised morphology in Williams et al. (2000).
Pelagodiscus atlanticus: Following table 6 (for Discinidae) in Williams et al. (2000).
Siphonobolus priscus: Noted in Siphonobolus by Havlicek (1982).
Terebratulina: “In modern terebratulides, the vascula media are subordinate to the lemniscate or pinnate vascula genitalia” – Williams et al. 1997 p417.
Tomteluva perturbata: Preservation not adequate to evaluate (Streng 2016).
Yuganotheca elegans: Based on the figures and sketches in Zhang et al. 2014 (and supplementary material), the mantle canals are interpreted as lateral, with no clear vascula media present.

[53] vascula terminalia

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[?]Haplophrentis carinatus[-]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[1]Micromitra[1]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[2]Botsfordia[2]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[2]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 53: Mantle canals: vascula terminalia

1: Exclusively marginal (peripheral)2: Directed peripherally and (intero)medially-: InapplicableTransformational character.

Presumed to be connected with setal follicles in life (Williams et al. 1998T). See Williams et al. (2000) for discussion.

Acanthotretella spinosa: Preservation not clear enough to score with certainty (Holmer & Caron 2006).
Alisina: Interomedial vascula terminalia not reported by Williams et al. (2000).
Askepasma toddense, Micromitra: Peripheral only (Williams et al. 1998T; Williams et al. 2000).
Botsfordia, Eoobolus: Following Williams et al. 1998T, appendix 2.
Glyptoria: Following appendix 2 in Williams et al. (1998T).
Heliomedusa orienta: Inferred from Jin & Wang (1992).
Kutorgina chengjiangensis, Salanygolina: Coded uncertain in appendix 2 in Williams et al. (1998T).
Lingula: Peripheral and medial for all Lingulata (Williams et al. 2000).
Lingulellotreta malongensis: Not described in Williams et al. (2000).
Lingulosacculus: Strong indication of medially directed vascula terminalia from vascula lateralia; see fig. 1.A1 in Balthasar & Butterfield 2009E.
Novocrania: Peripheral only (Williams et al. 2000, p.158).
Orthis: See schematics in Williams et al. (2000).
Pelagodiscus atlanticus: Following Lochkothele (Discinidae), fig. 43.4a in Williams et al. (2000).
Siphonobolus priscus: Not reported in Havlicek 1982 or Williams et al. 2000.
Terebratulina: Following idealised plectolophous terebratulid of Emig (1992).

4.8 Perioral tentacular apparatus

[54] Presence

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[0]Capitella[0]Canadia spinosa[1]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[1]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[1]Alisina[1]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 54: Perioral tentacular apparatus: Presence

0: Absent1: Present?: AmbiguousNeomorphic character.

The lophophore is a ring of tentacles that surrounds the mouth. Temereva (2017) suggests that true lophophores must also encompass the anus, which excludes the tentacular apparatus of entoprocts from the definition; as homology between the tentacular apparatuses of entoprocts and other lophophorates has often been assumed, we prefer to take a more inclusive stance and code the structures as potentially homologous.

It is unlikely that the tentacles of annelids and sipunculans correspond to the lophophore, yet homology is not inconceivable. In order that the tentacular apparatus of Haplophrentis can be compared with both organs without prejudice, we capture the presence of a tentacular apparatus in this very broad character, with arguments against homology reflected in separate transformation series.

Canadia spinosa, Serpula: Palps present (Parry & Caron, 2019).
Capitella: Palps absent (Parry & Caron, 2019).
Cotyledion tylodes: The tentacular crown (Zhang et al., 2013) is interpreted as a lophophore.
Dentalium: The scaphopod captacula is conceivably equivalent to the tentacular apparatus of other lophotrochozoans. It is developmentally pre-oral, and has tentatively been homologised with the pre-oral tentacles of Monoplacophora and Gastropoda (Steiner, 1992), though their musculature and late development suggests instead that they may derive from the molluscan foot, as do the arms of cephalopods (Wanninger & Haszprunar, 2002b).
Haplophrentis carinatus: Moysiuk et al. (2017).

[55] Origin

Character adds 3 to tree score (2 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[-]Capitella[1]Canadia spinosa[1]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[?]Amathia[2]Loxosomella[?]Namacalathus[3]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 55: Perioral tentacular apparatus: Origin

1: Prostomium (i.e. anterior of larval prototroch)2: Second pair of coelomic sacs, at metamorphosis3: Mid-trunk, prior to metamorphosis?: Ambiguous-: InapplicableTransformational character.

The tentacles of annelids and sipunculans originate from a dorsal pair of buds on the prostomium (Adrianov, Malakhov, & Maiorova, 2006), whereas the brachiopod lophophore arises from the second pair of coelomic sacs (Nielsen, 1991).

Canadia spinosa: Prostomial palps (Parry & Caron, 2019).
Dentalium: The captacula arise close to the mouth after metamorphosis (Wanninger & Haszprunar, 2002b), in a position not dissimilar from that of the phoronid tentacles (Santagata, 2004).
Flustra: The tentacles appear at metamorphosis, seemingly from below the corona (=prototroch) (Young, 2002).
Loxosomella: Arising after metamorphosis (Nielsen, 1971).

Novocrania: “At metamorphosis [….] the second pair of coelomic sacs develop small attachment areas at the edge of the dorsal valve and become the lophophore coelom” (Nielsen, 1991)

“The larval lobes are retained during the first steps of metamorphosis and are
subsequently remodeled to form the lophophore and other adult organs” – Altenburger, Wanninger, & Holmer (2013).
Phoronis: At the posterior of the head, at the late larval stage (Santagata, 2004).
Sipunculus: (Adrianov et al., 2006).
Terebratulina: Lophophore of Terebratalia arises post metamorphosis (Young, 2002); lophophore conceivably arising from vesicular bodies at base of apical lobe?

[56] Tentacle disposition

Character adds 4 to tree score (3 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[-]Capitella[-]Canadia spinosa[1]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[1]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 56: Perioral tentacular apparatus: Tentacle disposition

1: Single side2: Both sides?: Ambiguous-: InapplicableTransformational character.

Tentacles may occur along one or both sides of the axis of the lophophore arm (Carlson 1995).

Acanthotretella spinosa: Preservation insufficient to evaluate (Holmer & Caron 2006).
Alisina, Lingulosacculus, Longtancunella chengjiangensis: Preservation inadequate.
Amathia: Single side (Temereva & Kosevich, 2016).
Cotyledion tylodes: Tentacles seemingly occupy a single side of the lophophore (Zhang et al., 2013).
Dentalium: On rim of basal lobe only (Morton, 1959).
Flustra: Both sides (Schwaha & Wanninger, 2015; Shunkina, Zaytseva, Starunov, & Ostrovsky, 2015).
Heliomedusa orienta: “Each lophophoral arm bears a row of long, slender flexible tentacles” – Zhang et al. 2009.
Kutorgina chengjiangensis: Tentacles “cannot be confidently demonstrated in the available specimens.” – Zhang et al. 2007R.
Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina, Phoronis: Following coding for higher group in Carlson 1995, appendix 1, character 36.
Lingulellotreta malongensis: “The tentacles are clearly visible, and closely arranged in a single palisade” – Zhang et al. 2004N.
Loxosomella: Single side (Nielsen, 1966).
Sipunculus: Both sides in tentacle-breathers such as Themiste (Adrianov et al., 2006; Ruppert & Rice, 1995); only one side in Sipunculus (Adrianov et al., 2006; Ruppert & Rice, 1995).

[57] Tentacle rows per side in trocholophe stage

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 57: Perioral tentacular apparatus: Tentacle rows per side in trocholophe stage

0: No additional ablabial row1: Adlabial and ablabial rowNeomorphic character.

After Carlson (1995), character 37. Lophophore tentacles are commonly arranged into an ablabial and adlabial row, with ablabial tentacles sometimes added later in development.

Amathia: (Temereva & Kosevich, 2016).
Loxosomella, Flustra: Inapplicable.
Lingula, Pelagodiscus atlanticus, Terebratulina, Phoronis: Following coding for higher taxon in Carlson (1995), appendix 1, character 37.
Novocrania: Following coding for higher taxon in Carlson (1995), appendix 1, character 37. Also states in Williams et al. 2000, p. 158.

[58] Tentacle rows per side in post-trocholophe stage

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 58: Perioral tentacular apparatus: Tentacle rows per side in post-trocholophe stage

0: No additional ablabial row1: Adlabial and ablabial row?: AmbiguousNeomorphic character.

After Carlson (1995), character 37. Lophophore tentacles are commonly arranged into an ablabial and adlabial row, with ablabial tentacles sometimes added later in development (and thus interpreted as a neomorphic addition).

Acanthotretella spinosa: Preservation insufficient to evaluate (Holmer & Caron 2006).
Amathia: (Temereva & Kosevich, 2016).
Cotyledion tylodes: Additional row not evident (Zhang et al., 2013).
Heliomedusa orienta: “The lophophoral arms bear laterofrontal tentacles with a double row of cilia along their lateral edge, as in extant lingulid brachiopods” – Zhang et al. 2009.
Kutorgina chengjiangensis: Tentacles “cannot be confidently demonstrated in the available specimens.” – Zhang et al. 2007R.
Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina, Phoronis: Following coding for higher taxon in Carlson (1995), appendix 1, character 37.
Lingulellotreta malongensis: Single palisade (Zhang et al. 2004N).
Lingulosacculus: Preservation insufficient to evaluate.
Loxosomella: Nielsen (1966).
Yuganotheca elegans: “helical lophophore fringed with a single row of thick, widely spaced, parallel-sided and hollow tentacles” – Zhang et al. 2014.

[59] Median tentacle in early development

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 59: Perioral tentacular apparatus: Median tentacle in early development

0: Absent1: PresentNeomorphic character.

Following character 28 in Carlson 1995. Certain taxa exhibit a median tentacle early in development that is lost during ontogeny.

Namacalathus, Dailyatia, Acanthotretella spinosa, Alisina, Askepasma toddense, Micromitra, Antigonambonites planus, Clupeafumosus socialis, Coolinia pecten, Eccentrotheca, Gasconsia, Glyptoria, Heliomedusa orienta, Kutorgina chengjiangensis, Lingulosacculus, Lingulellotreta malongensis, Longtancunella chengjiangensis, Micrina, Mummpikia nuda, Nisusia sulcata, Orthis, Paterimitra, Salanygolina, Tomteluva perturbata, Yuganotheca elegans, Haplophrentis carinatus, Pedunculotheca diania: Lophophore ontogeny presently unknown.
Loxosomella: Nielsen (1966).

[60] Site of tentacle addition

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[-]Capitella[-]Canadia spinosa[1]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[?]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 60: Perioral tentacular apparatus: Site of tentacle addition

1: At two points located behind the mouth2: At the tip of each lophophore arm?: Ambiguous-: InapplicableTransformational character.

Following Temereva (2017).

Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina, Flustra, Amathia: Following Temereva (2017).
Phoronis: Following Temereva (2017) – though in larvae, tentacles are added at the tips of the developing lophophore.
Sipunculus: New branches added at each lateral extreme, behind mouth (Adrianov et al., 2006).

[61] Innervation

Character adds 3 to tree score (2 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[-]Capitella[1]Canadia spinosa[1]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[1]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 61: Perioral tentacular apparatus: Innervation

1: Extensions of a circumoral nerve ring2: Nerve rings within the tentacle ring itself?: Ambiguous-: InapplicableTransformational character.

Annelid tentacles are innervated by palp nerves (Orrhage & Müller, 2005); lophophores ancestrally contained a pair of nerve rings (Temereva, 2017).

Flustra, Amathia: Following Temereva (2017).
Canadia spinosa: Palp innervation – Parry & Caron (2019).
Dentalium: The captacula each bear an individual nerve fibre emanating from the cerebral ganglia, which is also associated with the circumoesophageal nerve ring (Sumner-Rooney et al., 2015), recalling the situation in annelids and sipunculans.
Loxosomella: Tentacle nerves originate laterally from the cerebral ganglion, branching three times and leading to a single nerve within each tentacle (Fuchs, Bright, Funch, & Wanninger, 2006).
Serpula: Orrhage & Müller (2005).
Sipunculus: Rice (1993).

[62] Inner nerve ring

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 62: Perioral tentacular apparatus: Inner nerve ring

0: Not reduced (whether present or absent due to absence of lophophore nerve rings)1: Reduced, weakly developed or absent in adults?: AmbiguousNeomorphic character.

Juvenile lophophorates exhibit two nerve rings in the tentacles; one of these rings is often reduced or lost at adulthood (Temereva, 2017).

Flustra, Amathia: Following Temereva (2017).
Lingula: Temereva & Kuzmina (2017).
Loxosomella: Nerves present in tentacles, but not forming rings (Fuchs et al., 2006).
Novocrania: Probably only a single ring is present, but only available illustrations are 19th century sketches (Lüter, 2016).
Phoronis: (Temereva, 2017).
Terebratulina: In Gryphus (Temereva & Kuzmina, 2017).

[63] Outer nerve ring

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 63: Perioral tentacular apparatus: Outer nerve ring

0: Not reduced (whether present or absent due to absence of lophophore nerve rings)1: Reduced, weakly developed or absent in adultsNeomorphic character.

Juvenile lophophorates exhibit two nerve rings in the tentacles; one of these rings is often reduced or lost at adulthood (Temereva, 2017).

Amathia: Following Temereva (2017); only one tentacle nerve ring evident in Temereva & Kosevich (2016).
Flustra: “Most species of bryozoans have only the inner” nerve ring – Temereva (2017).
Lingula, Terebratulina: Temereva (2017).
Loxosomella: Nerves present in tentacles, but not forming rings (Fuchs et al., 2006).
Novocrania: Probably only a single ring is present, but only available illustrations are 19th century sketches (Lüter, 2016).
Phoronis: Temereva & Kuzmina (2017).

[64] Musculature

Character adds 2 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[-]Capitella[?]Canadia spinosa[2]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[3]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[1]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 64: Perioral tentacular apparatus: Musculature

1: Outer main tentacle muscle; two pairs of inner longitudinal muscles2: Peripheral muscle fibres3: Core of longitudinal muscle fibres surrounded by circular muscles?: Ambiguous-: InapplicableTransformational character.

Dentalium: Six to eight elongate muscle cells in core (Shimek, 1988), surrounded by circular muscles (Byrum & Ruppert, 1994).
Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina: “Inner coelomic epithelium underlain by muscle fibers, or in the tentacles, myoepithelial cells.” – Williams et al. (1997).
Loxosomella: Outer main tentacle muscle; two pairs of inner longitudinal muscles (Fuchs et al., 2006).
Phoronis: (Pardos, Roldán, Benito, & Emig, 1991).
Serpula: Peripheral muscle fibres (Hanson, 1949).
Sipunculus: Peripheral to main tentacle cavity (Pilger, 1982).

[65] Forms closed loop

Character adds 6 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[-]Capitella[-]Canadia spinosa[2]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[2]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[2]Alisina[1]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[2]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[2]Acanthotretella spinosa[2]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 65: Perioral tentacular apparatus: Forms closed loop

1: Diverging laterally2: Closed loop?: Ambiguous-: InapplicableTransformational character.

Whereas the lophophore of crown-group brachiopods typically forms a closed loop, those of Haplophrentis and Heliomedusa diverge laterally (Moysiuk et al 2017).

Amathia: Ends of arms meet to form closed loop (Temereva & Kosevich, 2016).
Cotyledion tylodes: Tentacles form almost complete circular crown.
Lingulosacculus: Two diverging arms of the lophophore are preserved (Balthasar & Butterfield 2009E).
Longtancunella chengjiangensis: Two distinct, diverging arms reconstructed by Zhang et al. 2007A.
Loxosomella: Nielsen (1966).
Namacalathus: The existence of a lophophore is speculative.
Nisusia sulcata: No specimens of Nisusia preserve the lophophore.
Phoronis: Two lophophore arms rather than a single continuous loop, but with tips close together to form functional loop (Torrey, 1901).
Sipunculus, Serpula: Growing to encircle mouth in adults.

[66] Coiling direction

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[-]Capitella[-]Canadia spinosa[2]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[2]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[1]Alisina[1]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[2]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 66: Perioral tentacular apparatus: Coiling direction

1: Anteriad2: Posteriad-: InapplicableTransformational character.

The lophophore arms of Heliomedusa and Haplophrentis arch posteriad, rather than anteriad as in lingulids. See Zhang et al. 2009; Moysiuk et al. 2017.

Acanthotretella spinosa, Lingulellotreta malongensis: Arms proceed anteriad before recurving.
Flustra, Amathia: Bryozoan arms reach in anal (i.e. posterior) direction (Shunkina et al., 2015).
Cotyledion tylodes: Cannot establish without distinguishing gut from anus.
Loxosomella: Posterior (anal side) of lophophore has short stretch lacking tentacles.
Pelagodiscus atlanticus: “converging anteriorly and coiling anterior to the body cavity” – Zhang et al. 2009.
Phoronis: Coiling in direction of anus (i.e. posteriad).

[67] Adjustor muscle

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 67: Perioral tentacular apparatus: Adjustor muscle

0: Absent1: Present?: AmbiguousNeomorphic character.

Following character 55 in Carlson (1995). Not possible to code in most fossil taxa.

Namacalathus, Dailyatia, Acanthotretella spinosa, Alisina, Askepasma toddense, Micromitra, Antigonambonites planus, Clupeafumosus socialis, Coolinia pecten, Eccentrotheca, Gasconsia, Glyptoria, Heliomedusa orienta, Kutorgina chengjiangensis, Lingulosacculus, Lingulellotreta malongensis, Longtancunella chengjiangensis, Micrina, Mummpikia nuda, Nisusia sulcata, Orthis, Paterimitra, Salanygolina, Tomteluva perturbata, Yuganotheca elegans, Haplophrentis carinatus, Pedunculotheca diania: Preservation not adequate to evaluate presence or absence of this muscle.
Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina, Phoronis: Following coding for higher taxon in Carlson (1995), appendix 1, character 55.

4.9 Mouthparts: Buccal organ

[68] Buccal organ

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[1]Capitella[1]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 68: Mouthparts: Buccal organ

0: Absent1: Present?: AmbiguousNeomorphic character.

The buccal organ describes the structures that arise from the larval mouth region. This may include the foregut, which if eversible is termed a proboscis, and whose muscular regions are termed the pharynx (???).

Sipunculans express a buccal organ as larvae, which is lost after metamorphosis (???).

Novocrania, Lingula, Canadia spinosa, Serpula, Capitella: Following closest relative in Parry & Caron (2019).
Sipunculus: Present in larvae, following (???), fig. 83.

[69] Eversible

Character adds 0 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[2]Capitella[2]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 69: Mouthparts: Buccal organ: Eversible

2: Eversible-: InapplicableTransformational character.

Character 133 in Parry & Caron (2019).

[70] Papillae

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[1]Capitella[0]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 70: Mouthparts: Buccal organ: Papillae

0: Absent1: PresentNeomorphic character.

Character 134 in Parry & Caron (2019).

Sipunculus, Canadia spinosa, Capitella: Following Parry & Caron (2019).

4.10 Radula [71]

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[0]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 71: Radula

0: Absent1: PresentNeomorphic character.

Character 25 in Vinther et al. (2017). Any apparatus comprising multiple denticulate rows arranged serially in the sagittal plane is treated as potentially homologous with the molluscan radula.

Acaenoplax hayae: Seemingly absent (???).
Acanthotretella spinosa, Heliomedusa orienta, Kutorgina chengjiangensis, Lingulosacculus, Lingulellotreta malongensis, Longtancunella chengjiangensis, Yuganotheca elegans, Haplophrentis carinatus: No candidate observed despite exceptional preservation.
Kulindroplax perissokomos: No radula is preserved (Sutton et al., 2012).
Orthrozanclus: The radula has a high preservation potential in Wiwaxia, and is not evident only when it occurs at a different plane to the one that the fossil splits upon. As such, it is difficult to attribute the absence of a radula in Orthrozanclus to non-preservation.
Wiwaxia corrugata: 1.

[72] Extent

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[3]Wirenia[1]Chaetoderma[3]Tonicella[3]Mopalia[?]Polysacos vickersianum[3]Leptochiton[?]Glaphurochiton carbonarius[3]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[2]Wiwaxia corrugata[2]Odontogriphus omalus[3]Neopilina[?]Conocardium elongatum[3]Haliotis[?]Pojetaia runnegari[-]Mytilus[3]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[-]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[-]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[-]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 72: Radula: Extent

1: Single row of teeth2: Restricted: two to three tooth rows3: Extensive: myriad tooth rows-: InapplicableTransformational character.

Character 26 in Vinther et al. (2017). The radulae of Wiwaxia and Odontogriphus are conspicuously similar in their configuration.

Wirenia: Many rows in e.g. Plawenia (Amélie H. Scheltema, 2014).

[73] Subradular membrane

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 73: Radula: Subradular membrane

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 65 in Haszprunar (2000).

[74] Alary processes

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 74: Radula: Alary processes

0: Absent1: Present?: AmbiguousNeomorphic character.

A robust structure (alary process/hyaline shield) attached to the radula, with thickened margins, increasingly labile towards the rear, and constructed from the same material (chitin) as the radular teeth (Smith, 2012b).
Also referred to as a ‘hyaline shield’.

Odontogriphus omalus: Present (Smith, 2012b).
Wiwaxia corrugata: Inferred to be present (Smith, 2012b), but not observed, so coded as ambiguous.

[75] Bolster vesicles

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 75: Radula: Bolster vesicles

0: Absent1: Present?: AmbiguousNeomorphic character.

Hollow fluid-filled radula-supporting structures found in Polyplacophora and Monoplacophora (Katsuno & Sasaki, 2008).

[76] Subradular organ

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[0]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 76: Radula: Subradular organ

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 3g in Waller (1998); Character 58 in Haszprunar (2000).

[77] Heterodonty

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[1]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[2]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[-]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[-]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[-]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[-]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 77: Radula: Teeth: Heterodonty

1: Homodont: lateral teeth have identical morphology, if differing in size2: Heterodont: lateral teeth with different morphologies?: Ambiguous-: InapplicableTransformational character.

Character 29 in Vinther et al. (2017). Inapplicable if multiple lateral teeth are not present.

Calvapilosa kroegeri: Lateral and uncinical teeth, as in chitons (Vinther et al., 2017).
Haliotis: All teeth bear multiple cusps.

[78] Bending plane

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[2]Wiwaxia corrugata[2]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[-]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 78: Radula: Teeth: Bending plane

1: Stereoglossate2: Flexoglossate?: Ambiguous-: InapplicableTransformational character.

Character 60 in Ponder & Lindberg (1997); 2.20 in (???).

[79] More teeth per row in larger individuals

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[-]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[-]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[-]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[-]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[-]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[-]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 79: Radula: Teeth: More teeth per row in larger individuals

1: Ontogenetic increase in tooth number2: Number of teeth per row fixed throughout ontogeny?: Ambiguous-: InapplicableTransformational character.

Calvapilosa kroegeri: “exact number [of teeth in a row] is difficult to discern” (Vinther et al., 2017).

[80] Lateral tooth base

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[0]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 80: Radula: Teeth: Lateral tooth base

0: Absent1: Present?: AmbiguousNeomorphic character.

Presence of a distinct base in lateral teeth; see character 9 in Reynolds & Okusu (1999) and 15 in Steiner (1998).

[81] Lateral tooth head

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 81: Radula: Teeth: Lateral tooth head

0: Scarcely differentiated1: Prominently differentiated?: Ambiguous-: InapplicableNeomorphic character.

In polyplacophorans, the head of the lateral tooth is elaborate or clearly differentiated from the shaft ((???), character 8).

Oh dear! You included the inapplicable token in a neomorphic character!
That’s unlikely to be a good idea: see Brazeau, Smith & Guillerme (2018, Systematic Biology).
Unless you are very sure that you understand the consequences, you should mark the character as Transformational by setting State 0 to [Transformational character], or re-code:

  • Wirenia
  • Chaetoderma

[82] Apatite

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 82: Radula: Teeth: Apatite

0: Absent1: Present?: AmbiguousNeomorphic character.

Polyplacophoran teeth are reinforced with apatite (Haszprunar, 2000, character 69).

[83] Magnetite

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 83: Radula: Teeth: Magnetite

0: Absent1: Present?: AmbiguousNeomorphic character.

The tips of polyplacophoran teeth contain magnetite (Waller, 1998, character 4e).

4.11 Digestive tract

[84] Prominent pharynx

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[1]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[1]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 84: Digestive tract: Prominent pharynx

0: Absent1: Present?: AmbiguousNeomorphic character.

Hyoliths exhibit a prominent protrusible muscular pharynx at the base of the lophophore (Moysiuk et al. 2017). This is considered as potentially equivalent to the anterior projection of the visceral cavity in Heliomedusa, and, by extension, in Lingulosacculus and Lingulotreta.

Calvapilosa kroegeri: Not evident (Vinther et al., 2017), and interpreted as absent based on position of radula.
Canadia spinosa: The interpretation of a possible pharynx is ambiguous (???, ???).
Capitella: ‘Prominence’ is perhaps arguable in Capitella.
Eoobolus: Prominent extension of dorsal visceral platform (Balthasar 2009T).
Heliomedusa orienta: Corresponding to the “neck” of the vase-shaped visceral cavity reported by Zhang et al. 2009.
Lingulellotreta malongensis: An anterior projection of the visceral area is noted by Williams et al. (2000) and considered equivalent to that observed in Lingulosacculus (Balthasar & Butterfield 2009E).
Lingulosacculus: The prominent anterior extension of the visceral area noted by Balthasar & Butterfield (2009E) is considered as potentially homologous with that of Heliomedusa (Zhang et al. 2009) and, by extension, Haplophrentis (Moysiuk et al. 2017).
Sipunculus: Eversible pharynx (introvert).
Yuganotheca elegans: Possibly present, following interpretation of mouth (see fig. 2c, d in Zhang et al. 2014).

[85] Oesophageal folds

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 85: Digestive tract: Oesophageal folds

0: Absent1: Present?: AmbiguousNeomorphic character.

Following character 86 in Giribet & Wheeler (2002).

Phoronis: Ciliated ridge in oesophagus (Torrey, 1901).

[86] Oral sphincter

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[?]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 86: Digestive tract: Oral sphincter

0: Absent1: PresentNeomorphic character.

Character 133 in Grobe (2007).

Dentalium: Present, but secondarily reduced.

[87] Reduced

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 87: Digestive tract: Oral sphincter: Reduced

0: Not reduced1: Secondarily reduced?: AmbiguousNeomorphic character.

Character 133 in Grobe (2007).

[88] Locomotory cilia

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 88: Digestive tract: Foregut: Locomotory cilia

0: Absent1: PresentNeomorphic character.

Character 66 in Haszprunar (2000).

[89] Subdivisions

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 89: Digestive tract: Midgut: Subdivisions

0: Not subdivided1: Functional subdivisionsNeomorphic character.

The molluscan midgut is functionally subdivided into a sorting area (stomach), digestion area (midgut sac or gland), and transport tube (intestine). Characters 42 in Haszprunar (2000), 1.38 in von Salvini-Plawen & Steiner (1996).

Canadia spinosa: The gut is a straight tube with no obvious subdivision (Parry & Caron, 2019).
Odontogriphus omalus, Wiwaxia corrugata: Subdivided, presumably functionally, but with some ambiguity [1;Smith2014].

[90] Glands

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 90: Digestive tract: Midgut: Glands

0: Absent1: PresentNeomorphic character.

Characters 1.40, 2.30 and 4.59 in von Salvini-Plawen & Steiner (1996); 42 in Haszprunar (2000).

Odontogriphus omalus, Wiwaxia corrugata: Annex to midgut interpreted as a gland (Smith, 2012b).

4.12 Digestive tract: Anus

[91] Presence

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[0]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[0]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[0]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[0]Longtancunella chengjiangensis[?]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 91: Digestive tract: Anus: Presence

0: Anus present: through-gut1: Anus lost: digestive tract is blind sac?: AmbiguousNeomorphic character.

The digestive tract may either constitute a blind sac, or a through gut with anus. The loss of an anus is known to be derived within spiralia, so this character is treated as neomorphic.

Glyptoria: Scored according to familial level feature.
Kulindroplax perissokomos: Interpreted as possessing a through gut.
Kutorgina chengjiangensis: Although “the possibility of a blind ending may not be completely eliminated […] the weight of evidence […] leads us to reject the possibility of a blind-ending intestine” – Zhang et al. 2007R, p. 1399.

[92] Location

Character adds 8 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[2]Sipunculus[3]Wirenia[13]Chaetoderma[3]Tonicella[3]Mopalia[?]Polysacos vickersianum[3]Leptochiton[?]Glaphurochiton carbonarius[13]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[3]Neopilina[?]Conocardium elongatum[3]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[13]Kulindroplax perissokomos[?]Paramicrocornus[2]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[2]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[3]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[2]Cotyledion tylodes[?]Eccentrotheca[1]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[2]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[2]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[2]Acanthotretella spinosa[2]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 92: Digestive tract: Anus: Location

1: Straight gut with posterior anus2: Anus migrated posteriad to create U-shaped gut3: Anus opening to rear of pedal sole, causing slight U-shape to gut?: AmbiguousTransformational character.

“The relative position of the mouth and anus in the larvae of brachiopods and phoronids is similar: posterior anus and anterior mouth” – Williams et al. 2007, p. 2884
See also character 6 in Haszprunar & Wanninger (2008).

Chaetoderma: Coded as ambiguous (straight / rear of pedal sole) as the pedal sole is secondarily lost in the group.
Dentalium: The U-shaped gut of scaphopods arises by exaggeration of the dorsal surface, rather than migration of the anus (Steiner, 1992).
Kutorgina chengjiangensis: “Five specimens have an exceptionally preserved digestive tract, dorsally curved, with a putative dorso-terminal anus located near the proximal end of a pedicle” – Zhang et al. 2007R.
Terebratulina: “In rhynchonelliforms, the gut curves somewhat into a C-shape and the (blind) anus becomes posteroventral in position.” – Williams et al. 2007, p.
2884.

[93] Migration: Within ring of tentacles

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[2]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[2]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[1]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 93: Digestive tract: Anus: Migration: Within ring of tentacles

1: Not within ring of tentacles2: Anterior - within ring of feeding tentacles?: Ambiguous-: InapplicableTransformational character.

A migrated anus may be located laterally or within the lophophore ring (as in entoprocts).

Kutorgina chengjiangensis: “Presumed to terminate in a functional anus located near the proximal end of the pedicle.” – Zhang et al. 2007R.

[94] Migration: Position

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[3]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[2]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[4]Flustra[4]Amathia[-]Loxosomella[?]Namacalathus[3]Phoronis[?]Dailyatia[-]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[4]Terebratulina[?]Orthis[?]Alisina[2]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[3]Kutorgina chengjiangensis[3]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[3]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[2]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 94: Digestive tract: Anus: Migration: Position

2: Right3: Dorsally4: Ventrally?: Ambiguous-: InapplicableTransformational character.

If the anus is not within the ring of tentacles, in which direction is it oriented?

Flustra, Amathia: Anus remains on ventral surface. Arguably, rather than the anus migrating, the dorsal surface of the animal has become extended.
Dentalium: An alternative interpretation would be that the posterior of the scaphopod has been extended to generate the relatively anterior position of the originally ventral anus.
Haplophrentis carinatus: Opening to the right – see figures 1, 3, and extended data 5 in Moysiuk et al. (2017). The text states in error that the anus is to the left of the midline.
Kutorgina chengjiangensis: “Five specimens have an exceptionally preserved digestive tract, dorsally curved, with a putative dorso-terminal anus located near the proximal end of a pedicle” – Zhang et al. 2007R.
Lingula: “In the lingulids, the [intestine] follows an oblique course anteriorly to open at the anus on the right body wall.” – Williams et al. 1997, p. 89.
Lingulellotreta malongensis: “finally terminating in an anal opening on the right anterior body wall” (Zhang et al. 2007N, p.66).
Lingulosacculus: “This same arrangement occurs in L. nuda, with the looped dark line tracking the same course as the exceptionally preserved guts of Chengjiang lingulellotretids, including the median position of its posterior loop and the sharp right turn as it exits the posterior extension of the ventral valve” (Balthasar & Butterfield 2009E, p.310).
Longtancunella chengjiangensis: “The intestine extends posteriorly, and then turns right to continue as a tortuous strand, finally terminating at the latero-median position of the anterior body wall” – Zhang et al. 2007A.
Terebratulina: “In rhynchonelliforms, the gut curves somewhat into a C-shape and the (blind) anus becomes posteroventral in position.” – Williams et al. 2007, p.
2884.
Yuganotheca elegans: The identification of the “very poorly impressed possible anus at the lateral side of the anterior body wall” is not yet confident, so this character is coded as not presently available.

4.13 Sclerites

[95] Present in adult

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[1]Dentalium[1]Pelagiella[1]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[1]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[1]Dailyatia[1]Cotyledion tylodes[1]Eccentrotheca[1]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 95: Sclerites: Present in adult

0: Absent1: PresentNeomorphic character.

Plate-like (wider than tall) skeletal elements, whether mineralized or non-mineralized. Corresponds to character 8 in Vinther et al. (2017).
The definition deliberately excludes setae (which are taller than wide).

Tonicella, Dentalium: Molluscan valves are treated as potential homologues of brachiopod valves.
Halkieria evangelista: Halkieriid sclerites are interpreted as potentially homologous with those of Dailyatia and hence the brachiopods (Zhao et al., 2017).
Namacalathus: The mineralized endoskeleton of Namacalathus is not interpreted as a sclerite.
Serpula: Annelid setae are not considered to represent potential homologues with the brachiopod shell.
Sipunculus: Hooks are present, though the absence of chitin or microvillar impressions indicates that they are not homologous with those of other lophotrochozoans.
Wiwaxia corrugata: The scales of Wiwaxia are treated as homologous with the chaetae of annelids and brachiopods (Butterfield, 1990; Smith, 2014; Zhang et al., 2015), rather than brachiopod shell.
Yilingia spiciformis: The plate-like structures on the dorsal surface of Yilingia (Chen et al., 2019) are plausibly interpreted as non-mineralized sclerites.

[96] Periodically shed and replaced

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[1]Dentalium[1]Pelagiella[1]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[2]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[1]Namacalathus[-]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[2]Orthrozanclus[?]Micromitra[1]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[1]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[1]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 96: Sclerites: Periodically shed and replaced

1: Retained 2: Periodically shed and replaced?: Ambiguous-: InapplicableTransformational character.

Certain taxa periodically slough and replace some of their individual sclerites during growth. Others continue to add to sclerites by marginal accretion throughout life.

Acaenoplax hayae: Valves grown by marginal accretion (???).
Orthrozanclus: Inferred by comparison with Halkieria.
Paterimitra: Larval shell present at tip of sclerites (???), indicating retention.
Siphogonuchites multa: Retained as set in shell matrix (Bengtson, 1992).
Yilingia spiciformis: The broad size and interlocking nature of sclerites suggests that they are not periodically shed and replaced, consistent with the absence of any specimens lacking sclerites.

[97] Prominent major valves

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[1]Dentalium[1]Pelagiella[1]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[1]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[1]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 97: Sclerites: Prominent major valves

0: Scleritomous: without differentiated dorsal valve1: Scleritome dominated by prominent dorsal valve (and, optionally, accompanying ventral or posterior valves)Neomorphic character.

Equivalent to “Sclerites: Bivalved” in H.-J. Sun et al. (2018), rephrased to reflect the variation in the number of conceivably homologous ‘major’ shell plates in Aculifera.

A differentiated ventral or posterior valve may be present in addition to a prominent anterior/dorsal valve, corresponding to the ‘head valve’ of chitons or the dorsal valve of brachiopods.

Tonicella: As larvae, polyplacophorans exhibit an anterior and a posterior shell field (Wanninger & Haszprunar, 2002a); subsequent subdivision of the posterior field gives rise to the posterior seven valves. Tonicella is thus tentatively coded as ‘bivalved’ to reflect the potential (if perhaps unlikely) homology with the paired elements of brachiopods.

[98] Reduced

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[1]Polysacos vickersianum[2]Leptochiton[2]Glaphurochiton carbonarius[2]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[2]Neopilina[2]Conocardium elongatum[2]Haliotis[2]Pojetaia runnegari[2]Mytilus[2]Dentalium[2]Pelagiella[?]Siphogonuchites multa[2]Acaenoplax hayae[2]Kulindroplax perissokomos[2]Paramicrocornus[1]Haplophrentis carinatus[2]Cupitheca holocyclata[2]Bactrotheca[2]Pedunculotheca diania[1]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[1]Dailyatia[1]Cotyledion tylodes[1]Eccentrotheca[1]Orthrozanclus[2]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[2]Yuganotheca elegans[2]Tomteluva perturbata[2]Salanygolina[2]Terebratulina[2]Orthis[2]Alisina[2]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[2]Novocrania[2]Craniops[2]Ussunia[2]Heliomedusa orienta[2]Gasconsia[2]Pelagodiscus atlanticus[2]Mickwitzia muralensis[2]Mummpikia nuda[2]Botsfordia[2]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[2]Siphonobolus priscus[2]Acanthotretella spinosa[2]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[1]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 98: Sclerites: Accessory sclerites: Reduced

1: Accessory sclerites not reduced2: Accessory sclerites absent: primary valves only?: Ambiguous-: InapplicableTransformational character.

Taxa in the bivalved condition may retain sclerites as small additional elements, such as the L-elements of Paterimitra (Skovsted et al. 2015). Hyolithid helens are coded as potentially homologous to these elements (following Moysiuk et al., 2017).

This character is treated as neomorphic, with accessory sclerites ancestrally present, recognizing the likely origin of brachiozoans (and Lophotrochozoans more generally) from a scleritomous organism.

Acaenoplax hayae, Kulindroplax perissokomos: The spines are interpreted as homologous with the girdle elements of polyplacophorans, i.e. as setae.
Cupitheca holocyclata: Helens never observed and considered absent (Skovsted et al., 2016).
Dentalium: The scaphopod valve arises posterior of the prototroch and is thus homologous with the posterior valves of Chiton, assuming that molluscan shell fields are homologous features.
Paramicrocornus: Helens absent, with no possible insertion point (Z.-L. Zhang et al., 2018).
Paterimitra: L-sclerites (Skovsted et al. 2009T).
Polysacos vickersianum: The annulus of spines is considered to represent accessory sclerites homologous to the main valves; see discussion under ‘adult setae’.
Siphogonuchites multa: It is possible that two shell morphs exist and belonged to the same individuals; or that other aggregations of spicules represent additional shell fields (Bengtson, 1992; Conway Morris & Chapman, 1996).
Tonicella: The intermediate shell plates arise by subdivision of the posterior shell field (Wanninger & Haszprunar, 2002a), and are thus treated as equivalent to the posterior valve rather than as distinct elements.
The girdle elements are homologous with annelid chaetae / brachiopod setae (Leise & Cloney, 1982), rather than sclerites.

[99] Arrangement

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[1]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[1]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[1]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[1]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 99: Sclerites: Accessory sclerites: Arrangement

0: Single field1: Peripheral and medial fields with distinct sclerite arrangementsNeomorphic character.

Following Zhao et al. (2017), and reflecting character 5 in Vinther et al. (2017).

Dailyatia: Following the reconstruction of Skovsted, Betts, Topper, & Brock (2015).
Yilingia spiciformis: Distinct dorsal and dorsolateral fields, albeit in phase (Chen et al., 2019).

[100] Symmetry

Character adds 7 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[1]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[1]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[1]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[1]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[1]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[1]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 100: Sclerites: Accessory sclerites: Symmetry

0: Dextral and sinistral forms absent1: Occuring in dextral and sinistral forms?: AmbiguousNeomorphic character.

Following Zhao et al. (2017).

Eccentrotheca: Skovsted, Brock, Paterson, Holmer, & Budd (2008).
Haplophrentis carinatus: Helens are symmetrical.
Yilingia spiciformis: Chen et al. (2019).

[101] Side slope shape

Character adds 0 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[2]Polysacos vickersianum[2]Leptochiton[2]Glaphurochiton carbonarius[2]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[2]Neopilina[2]Conocardium elongatum[2]Haliotis[2]Pojetaia runnegari[2]Mytilus[?]Dentalium[2]Pelagiella[2]Siphogonuchites multa[2]Acaenoplax hayae[2]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[2]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[2]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[2]Paterimitra[2]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 101: Sclerites: Prominent major valves: Side slope shape

2: Convex-: InapplicableTransformational character.

Following character 12 in (???).

Acaenoplax hayae: Convex (???).
Calvapilosa kroegeri: Seemingly convex (Vinther et al., 2017).
Haliotis: Auzoux-Bordenave et al. (2010).
Halkieria evangelista: Convex (???).
Kulindroplax perissokomos: “Weakly convex” (Sutton et al., 2012).
Novocrania: Convex (???).
Orthrozanclus: Convex (Conway Morris & Caron, 2007).
Pelagodiscus atlanticus: (???).
Phthipodochiton thraivensis: “Side slopes slightly convex, rounded” (Sutton & Sigwart, 2012).
Terebratulina: (???).
Tonicella: Gently convex (Connors et al., 2012).
Yilingia spiciformis: Seemingly convex in thin section (Chen et al., 2019).

[102] Additional major valves

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[1]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[1]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 102: Sclerites: Prominent major valves: Additional major valves

0: Absent1: Present?: AmbiguousNeomorphic character.

To reflect the single valve present in Orthrozanclus and the conceivable homology between the tail valve of Halkieria and the ventral valve of brachiopods.

Acaenoplax hayae: Eight valves, including V7v and V7d as separate valves.
Dailyatia: Absent – coded as lacking a prominent dorsal valve.
Leptochiton: Kaas (1981).
Paterimitra: The S2 sclerite (???) is treated as a prominent valve by virtue of its close affiliation with the S1 sclerite.
Pojetaia runnegari: The two valves are considered to be a single valve, separated along the midline and joined by the ligament, following the conventional interpretation of rostroconchs.
Yilingia spiciformis: No major valves (Chen et al., 2019).

[103] Additional valves: Nature

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[2]Polysacos vickersianum[2]Leptochiton[2]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[2]Acaenoplax hayae[2]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 103: Sclerites: Prominent major valves: Additional valves: Nature

1: Prominent ventral valve2: One or more valves posterior to ‘’head plate’’?: Ambiguous-: InapplicableTransformational character.

The ventral valve of brachiopods is unlikely to be equivalent to the tail valve of Halkieria or chitons.

Paterimitra: The S2 valve is treated as ventral valve as it is associated with a likely pedicle opening.

[104] Serially repeated

Character adds 1 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[2]Polysacos vickersianum[2]Leptochiton[2]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[2]Acaenoplax hayae[2]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[1]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 104: Sclerites: Posterior valves: Serially repeated

1: Single valve2: Serially repeated valves-: InapplicableTransformational character.

[105] Number

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[1]Polysacos vickersianum[2]Leptochiton[2]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[2]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 105: Sclerites: Posterior valves: Number

1: Six (including tail valve)2: Seven-: InapplicableTransformational character.

Vinther et al. (2017) (character 19) report five intermediate shell fields in Kulindroplax, Acaenoplax, multiplacophorans, and the larvae of Chaetoderma.

Acaenoplax hayae: Seven, counting V7d as separate from V7v (???).
Glaphurochiton carbonarius: Hoare & Mapes (1986).
Kulindroplax perissokomos: Seven valves including head and tail (Sutton et al., 2012).
Phthipodochiton thraivensis: Eight valves in total (Sutton & Sigwart, 2012).

[106] Apophyses

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 106: Sclerites: Posterior valves: Apophyses

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 31 in Vinther et al. (2017). Sutural laminae or apophyses are teeth that articulate adjacent shell plates in many polyplacophorans.

Acaenoplax hayae: (???).
Glaphurochiton carbonarius: Following Vinther et al. (2017).
Mopalia, Tonicella, Leptochiton: Present in modern chitons (Vinther et al., 2017).
Polysacos vickersianum: Present in multiplacophorans (Vinther et al., 2017).

[107] Jugal ridges

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 107: Sclerites: Posterior valves: Jugal ridges

0: Absent1: Present?: AmbiguousNeomorphic character.

A jugal ridge is a medial longitudinal ridge. Following character 13 in (???).

Acaenoplax hayae: (???).
Glaphurochiton carbonarius: Hoare & Mapes (1986).
Kulindroplax perissokomos: Sharp jugal angle but no ridge.

[108] Insertion plates

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 108: Sclerites: Posterior valves: Insertion plates

0: Absent1: PresentNeomorphic character.

Character 32 in Vinther et al. (2017)
“In the majority of recent chitons the articulamentum may form extensions beyond the margin of the tegmentum. These extensions, called insertion plates, occur on the lateral margins of intermediate valves, on the anterior margin of the head valve and posteriorly on the tail valve” (Schwabe, 2010).

Acaenoplax hayae: (???).
Glaphurochiton carbonarius: Following Vinther et al. (2017).
Polysacos vickersianum: Present in multiplacophorans (Vinther et al., 2017).
Tonicella: Present (Connors et al., 2012).

[109] Insertion plates: Slit

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 109: Sclerites: Posterior valves: Insertion plates: Slit

0: Absent1: PresentNeomorphic character.

Character 33 in Vinther et al. (2017).
“The distal edge of the insertion plates may be slitted or solid in different taxa. The bridges between the slits (or incisions) are called teeth and may either be smooth at their outside, roughened, or even strongly pectinate.” (Schwabe, 2010).

Mopalia: (???).
Polysacos vickersianum: Following Vinther et al. (2017).
Tonicella: Slits with slit rays (Connors et al., 2012).

[110] Insertion plates: Slit: Nature

Character adds 0 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 110: Sclerites: Posterior valves: Insertion plates: Slit: Nature

2: Single slit-: InapplicableTransformational character.

Character 34 in Vinther et al. (2017).

Mopalia: (???).
Tonicella: Coded following Vinther et al. (2017).

[111] Insertion plates: Pectinate

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[2]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 111: Sclerites: Posterior valves: Insertion plates: Pectinate

1: Not pectinate2: Pectinate-: InapplicableTransformational character.

Character 35 in Vinther et al. (2017).

Mopalia: (???).
Polysacos vickersianum: Following Vinther et al. (2017).
Tonicella: Coded following Vinther et al. (2017).

[112] Differentiated intermediate shell fields

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[1]Mopalia[2]Polysacos vickersianum[1]Leptochiton[2]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[2]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 112: Sclerites: Posterior valves: Differentiated intermediate shell fields

1: Intermediate shell fields homonomous2: Intermediate shell fields distinct from one another?: Ambiguous-: InapplicableTransformational character.

Following character 17 in Vinther et al. (2017), itself derived from character 7 in (???). A satisfactory definition for this character is not available; it is here taken to mean “intermediate shell fields are differentiated from one another”, rather than “differentiated from the head/tail valves” or “spatially non-overlapping”.

Acaenoplax hayae: (???).
Glaphurochiton carbonarius: Following Vinther et al. (2017).
Leptochiton: Kaas (1981).
Mopalia: (???).
Phthipodochiton thraivensis: No differentiated plate II, as in many polyplacophorans (Sutton & Sigwart, 2012).
Tonicella: Small degree of morphological differentiation (Connors et al., 2012).

[113] Laterally divided shell fields

Character adds 2 to tree score (1 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[2]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[1]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 113: Sclerites: Posterior valves: Laterally divided shell fields

1: Undifferentiated: Each shell field comprises a single plate2: Differentiated into multiple plates-: InapplicableTransformational character.

Per character 18 in Vinther et al. (2017), the intermediate shell fields of multiplacophorans comprise multiple plates.

Acaenoplax hayae: (???).
Mopalia: (???).

[114] Hinge line shape

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[1]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[1]Orthrozanclus[2]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[2]Salanygolina[1]Terebratulina[2]Orthis[2]Alisina[2]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[12]Kutorgina chengjiangensis[2]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[2]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 114: Sclerites: Bivalved: Hinge line shape

1: Astrophic2: Strophic?: Ambiguous-: InapplicableTransformational character.

Askepasma toddense, Micromitra, Glyptoria: Coded as strophic in Williams et al (1998T).
Botsfordia: Coded as dissociated in Williams et al. (1998T), appendix 2.
Craniops: Astrophic: rounded posterior margin (see fig. 91 in Williams et al. 2000).
Cupitheca holocyclata: (Skovsted et al., 2016).
Eoobolus: Coded as astrophic in Williams et al (1998T); see also Balthasar (2009).
Gasconsia: The straight posterior margin of Gasconsia contributes to an overall resemblance with the Chileids (Holmer, Popov, & Bassett, 2014).
Halkieria evangelista, Mickwitzia muralensis: Non-strophic.
Kutorgina chengjiangensis: Williams et al. (1998b) (appendix 2) and Williams et al. (2000) (p. 208) consider the hinge of Kutorgina to be stropic, whereas Bassett et al (2001) argue for an astropic interpretation – whilst noting that the arrangement is prominently different from other astrophic taxa. We therefore code this taxon as ambiguous.
Longtancunella chengjiangensis: “Longtancunella has an oval to subcircular shell with a very short strophic hinge line” – Zhang et al. 2011T.
Micrina: Non-strophic: see Holmer et al 2008.
Nisusia sulcata: “The strophic, articulated shells of the Kutorginata rotated on simple hinge mechanisms that are different from those of other rhynchonelliforms” (Williams et al. 2000, p. 208).
Novocrania: Craniides have a strophic posterior valve edge (Williams et al. 2007, table 39 on p. 2853): Novocrania’s “dorsal posterior margin” is “straight” (Williams et al. 2000, p. 171).
Salanygolina: Coded as strophic in Williams et al (1998T); see Holmer, Pettersson Stolk, Skovsted, Balthasar, & Popov (2009).
Tomteluva perturbata: “Tomteluvid taxa all have a strongly ventribiconvex, astrophic shell with a unisulcate commissure” – Streng et al. 2016, p5.
Tonicella: A linear hinge articulation does not exist between valves 1 and 2; nor would it exist between valves 1 and 8 were these adjacent (Connors et al., 2012).
Yuganotheca elegans: Not evident from fossil material; the possibility of a short strophic hinge line (as in Longtancunella) is difficult to discount.

[115] Enclosing filtration chamber

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 115: Sclerites: Bivalved: Enclosing filtration chamber

0: No filtration chamber, or open chamber1: Shells close to form enclosed filtration chamber?: AmbiguousNeomorphic character.

In crown-group brachiopods, the two primary shells close to form an enclosed filtration chamber. Further down the stem, taxa such as Micrina do not.

[116] Commissure: Exact correspondence of valve margins

Character adds 3 to tree score (2 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[12]Cupitheca holocyclata[2]Bactrotheca[2]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 116: Sclerites: Bivalved: Commissure: Exact correspondence of valve margins

1: Margins align exactly when valves closed2: Margins of different shape or size?: Ambiguous-: InapplicableTransformational character.

Orthothecid hyoliths can retract their operculum into their conical shell, in contrast to most other taxa, where the valves align exactly when they are closed, save perhaps for a pedicle notch or, in the case of hyolithids, depressions that allow the helens to protrude. Refers only to two prominent valves, not to additional sclerites of e.g. Eccentrothca.

Bactrotheca: Operculum interpreted as sitting inside conical shell (Marek, 1976).
Cupitheca holocyclata: “The width range of opercula matches that of the apertures” (H.-J. Sun et al., 2018), but it is not possible to determine whether this is an exact match or whether the operculum may have been slightly smaller, and hence retractable, as anticipated for orthothecids.
Paramicrocornus: Articulated specimens unknown.

[117] Commissure: Sulcate

Character adds 4 to tree score (2 with Fitch)2 additional regions (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[1]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[1]Micromitra[12]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[2]Tomteluva perturbata[1]Salanygolina[12]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[2]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[-]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 117: Sclerites: Bivalved: Commissure: Sulcate

1: Rectimarginate2: Uniplicate-: AmbiguousTransformational character.

The anterior commissure can be rectimarginate (i.e. straight), uniplicate (i.e. median sulcus in ventral valve), or sulcate (with median sulcus in dorsal valve).

Askepasma toddense: Coded as rectimarginate in Williams et al. (1998b), though note that the “ventral valve weakly to moderately sulcate” (Topper et al. 2013T); a similar description is provided by Williams et al. (2000). Coded as ambiguous for these two states accordingly.
Micromitra, Glyptoria, Kutorgina chengjiangensis, Salanygolina: Following appendix 2 in Williams et al. (1998T).
Terebratulina: “Anterior commissure rectimarginate to uniplicate” – uniplicate in fig. 1425.1c of Williams et al. (2006).

[118] Commissure: Circular

Character adds 9 to tree score (7 with Fitch)2 additional regions (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[2]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[2]Paramicrocornus[2]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[2]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[2]Yuganotheca elegans[1]Tomteluva perturbata[2]Salanygolina[1]Terebratulina[1]Orthis[2]Alisina[2]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[2]Mummpikia nuda[2]Botsfordia[2]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 118: Sclerites: Bivalved: Commissure: Circular

1: Continuous round outline with no corners (save at the hinge); 2: Lateral margins linear?: Ambiguous-: InapplicableTransformational character.

Shape of the commissure in plan view, ignoring any deflection arising due to articulation at the hinge (e.g. delthyrium/notothyrium). This character seeks to discriminate the essentially conical ‘conchs’ of orthothecid hyoliths from the polygonal ‘conchs’ of hyolithids. Triangular and oblong outlines are not distinguished, as this is not entirely independent of the strophic/astrophic nature of the hinge.

Acanthotretella spinosa: Round (Holmer & Caron, 2006).
Alisina: Sub-triangular (Zhang, Holmer, Popov, & Shu, 2011).
Askepasma toddense, Antigonambonites planus, Coolinia pecten, Glyptoria, Nisusia sulcata: Lateral margins subparallel, cf. Askepasma (Williams et al., 2000).
Bactrotheca: Operculum essentially round, though becoming flattened on functionally ventral surface (Dzik, 1980).
Botsfordia: Diverging lateral margins (G.-X. Li & Holmer, 2004).
Clupeafumosus socialis: Round (Topper, Harper, & Ahlberg, 2013).
Craniops, Siphonobolus priscus: Round (Williams et al., 2000).
Dentalium: Opening essentially polygonal (a rolled rectangle).
Eoobolus: Essentially triangular (Williams et al., 2000).
Gasconsia: Round, hinge notwithstanding (Hanken & Harper, 1985).
Halkieria evangelista: Anterior shell essentially triangular.
Kutorgina chengjiangensis: Lateral margins subparallel (Williams et al., 2000, fig. 125); clear angular corners in K. chengjiangensis (Holmer et al., 2018b).
Lingulellotreta malongensis: Linear, diverging lateral margins (Zhang et al., 2007a).
Lingulosacculus: Round (Balthasar & Butterfield, 2009).
Mickwitzia muralensis: Round (Balthasar, 2004).
Micrina: Mitral valve aperture essentially round (Holmer et al., 2008).
Micromitra: Lateral margins subparallel, cf. Askepasma (Robson & Pratt, 2001).
Mummpikia nuda: Polygonal (Balthasar, 2008).
Novocrania, Pelagodiscus atlanticus: Essentially round (Williams et al., 2000).
Orthis: Essentially round, hinge notwithstanding (Williams et al., 2000, fig. 523).
Paramicrocornus: Polygonal (Z.-L. Zhang et al., 2018).
Paterimitra: Broad posterior sinus: not directly comparable with brachiopod condition.
Salanygolina: Short subparallel stretch of lateral sides of adult shell (Holmer et al., 2009, fig. 1a), recalling D-shaped profile of Askepasma.
Tomteluva perturbata: Essentially round, sulcus notwithstanding (Streng, Butler, Peel, Garwood, & Caron, 2016).
Ussunia: Essentially round, hinge notwithstanding (Williams et al., 2000).
Yuganotheca elegans: Polygonal (Zhang et al., 2014).

[119] Commissure: Lateral margins

Character adds 7 to tree score (5 with Fitch)4 additional regions (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[1]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[2]Paramicrocornus[2]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[2]Yuganotheca elegans[-]Tomteluva perturbata[1]Salanygolina[2]Terebratulina[-]Orthis[2]Alisina[2]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[-]Pelagodiscus atlanticus[-]Mickwitzia muralensis[2]Mummpikia nuda[2]Botsfordia[2]Eoobolus[2]Lingulellotreta malongensis[3]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[-]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 119: Sclerites: Bivalved: Commissure: Lateral margins

1: Subparallel2: Diverging3: Initially diverging, becoming subparallel?: Ambiguous-: InapplicableTransformational character.

If lateral margins are linear, are the subparallel (i.e. commissure profile oblong, with long hinge) or diverging (i.e. commissure profile triangular, with short hinge)?

[120] Apophyses

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[1]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 120: Sclerites: Bivalved: Apophyses

0: Absent1: PresentNeomorphic character.

Micrina, like many brachiopods, bears tooth-like structures or processes that articulate the two primary valves. Caution must be applied before taxa are coded as “absent”, as teeth can be subtle and may be overlooked.

Alisina: “Strophic articulation with paired, ventral denticles, composed of secondary shell” – definition of family Trematobolidae in Williams et al. 2000.
Clupeafumosus socialis: No articulating processes evident or reported by Topper et al. (2013R).
Gasconsia: “Articulatory structure comprising ventral cardinal socket and dorsal hinge plate […] The shape of the shell probably correlates strongly with the unique type of articulation, which consists of a dorsal hinge plate that fits tightly into a cardinal socket in the ventral valve, with a concave homeodeltidium in the center of the ventral interarea” – Williams et al. 2000, p.184, concerning order Trimerellida.
Kutorgina chengjiangensis, Nisusia sulcata: Kutorginata don’t have teeth or dental sockets, but their shells are articulated by “two triangular plates formed by dorsal interarea, bearing oblique ridges on the inner sides” (Williams et al 2000, p. 211); this simple hinge mechanism is different from other rhynchonelliforms (Williams et al. 2000, p.208), but serves an equivalent purpose and is thus potentially homologous. We thus code kutorginids as present, using a subsequent character to capture difference in tooth morphology.
Mickwitzia muralensis: Not reported by or evident in Balthasar (2004).
Mummpikia nuda: No articulation structures are evident; instead, the propareas are rotated inwards (Balthasar 2008). The definition of Family Obolellidae in Williams et al. (2000) notes that articulation may be lacking or vestigial in the group.
Tomteluva perturbata: Tomteluvids […] lack articulation structures such as teeth and sockets (Streng et al. 2016).
Tonicella: The sutural laminae correspond in function and position to brachiopod apophyses (Connors et al., 2012), and so are coded as potentially homologous.
Ussunia: “articulatory structures poorly developed” – Williams et al. 2000, p. 192.

[121] Apophyses: Morphology

Character adds 6 to tree score (4 with Fitch)2 additional regions (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[-]Micromitra[-]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[-]Tomteluva perturbata[-]Salanygolina[2]Terebratulina[1]Orthis[2]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[3]Nisusia sulcata[3]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[-]Pelagodiscus atlanticus[-]Mickwitzia muralensis[-]Mummpikia nuda[-]Botsfordia[-]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[1]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 121: Sclerites: Bivalved: Apophyses: Morphology

1: Deltidiodont2: Cyrtomatodont3: Pseudodont?: Ambiguous-: InapplicableTransformational character.

Deltidiodont teeth are simple hinge teeth developed by the distal accretion of secondary shell; Cyrtomatodont teeth are knoblike or hook-shaped hinge teeth developed by differential secretion and resorption of the secondary shell (fig. 322 in Williams et al 1997).

Kutorginata (here represented by Kutorgina and Nisusia) don’t have teeth (apophyses) or dental sockets, but their shells are articulated by “two triangular plates formed by dorsal interarea, bearing oblique ridges on the inner sides” (Williams et al 2000, p. 211); this simple hinge mechanism is different from other rhynchonelliforms (Williams et al. 2000, p.208; table 13 character 30), and is described as a “pseudodont articulation” (Holmer et al. 2018E).

Antigonambonites planus, Glyptoria: Coded as deltidiodont in Benedetto (2009).
Kutorgina chengjiangensis: “Articulation characterized by two triangular plates formed by dorsal interarea, bearing oblique ridges on the inner sides” – Williams et al 2000, p. 211.
Micrina: The simple knob-like teeth of Micrina show no evidence of resprobtion or the hook-like shape that characterises Cyrtomatodont teeth.
Nisusia sulcata: The ‘teeth’ are formed by the distal lateral extensions from the ventral
pseudodeltidium fitting into the ‘sockets’ on the inner side of the dorsal interarea (Holmer et al. 2018E). (Coded as “deltidiodont teeth absent” in Benedetto (2009).).
Orthis: Coded as deltidiodont (in Eoorthis) in Benedetto (2009).
Terebratulina: Cyrtomatodont – see fig. 322 in Williams et al (2000).
Tonicella: Chiton apophyses (sutural laminae) are accretions deriving from the ventral shell layer of the intermediate and tail valves (Schwabe, 2010), so correspond to the deltidiodont situation in brachiopods.

[122] Apophyses: Dental plates

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[1]Orthis[0]Alisina[?]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 122: Sclerites: Bivalved: Apophyses: Dental plates

0: Absent1: PresentNeomorphic character.

Williams et al. 1997 (p362) write: “Teeth […] are commonly supported by a pair of variably disposed plates also built up exclusively of secondary shell and known as dental plates (Fig. 323.1, 323.3).”

Dewing (2001) elaborates: “Dental plates are near-vertical, narrow sheets of shell tissue between the anteromedian edge of the teeth and floor of the ventral valve. They are a composite structure, resulting from the growth of teeth over the ridge that bounds the ventral-valve muscle field.”

Williams et al. 2000 (p.201) write: “The denticles lack supporting structures in all Obolellida, but in Naukatida they are supported by an arcuate plate below the
interarea, the anterise (Fig. 119.3a)”.

The anterise is conceivably homologous with the dental plates, thus the presence of either is coded “present” for this character.

Antigonambonites planus: Coded as present (well developed) in Benedetto (2009).
Coolinia pecten: Coded as present following Dewing (2001), who seems to use the term Strophomenoids to encompass Coolinia, and attests to the presence of dental plates.
Gasconsia: Coded ambiguous to reflect the possibility that the hinge plate in trimerellids is homologous to the dental plates of other taxa, and has replaced the teeth themselves as the primary articulatory mechanism (see Williams et al. 2000, p. 184, for details of the articulation).
Glyptoria, Nisusia sulcata: Coded as absent in Benedetto (2009).
Orthis: Coded as present (short and recessive, in Eoorthis) in Benedetto (2009).

[123] Sockets

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[1]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 123: Sclerites: Bivalved: Sockets

0: Absent1: PresentNeomorphic character.

Simplified from Bassett et al. (2001) character 16.
This character is independent of apophyses, as several taxa bear sockets without corresponding teeth; the function of these sockets is unknown.
See figs 323ff in Williams et al. (1997).

Alisina: “bearing sockets, bounded by low ridges” – Williams et al. 2000.
Antigonambonites planus: Coded as present in Benedetto (2009).
Gasconsia: “Articulatory structure comprising ventral cardinal socket and dorsal hinge plate” – Williams et al. 2000, p. 184.
Glyptoria, Nisusia sulcata: Coded as absent in Benedetto (2009).
Mickwitzia muralensis: Not reported by or evident in Balthasar (2004).
Tomteluva perturbata: Tomteluvids […] lack articulation structures such as teeth and sockets (Streng et al. 2016).
Ussunia: Following table 15 in Williams et al. 2000.

[124] Socket ridges

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 124: Sclerites: Bivalved: Socket ridges

0: Absent1: PresentNeomorphic character.

After Bassett et al. (2001) character 17. May be difficult to distinguish from a brachiophore (see Fig 323 in Williams et al 1997), so the two structures are not distinguished here.

Alisina: “bearing sockets, bounded by low ridges” – Williams et al. 2000.
Antigonambonites planus: Coded as present in Benedetto (2009).
Glyptoria, Nisusia sulcata: Coded as absent in Benedetto (2009).
Tomteluva perturbata: Tomteluvids […] lack articulation structures such as teeth and sockets (Streng et al. 2016).

[125] Muscle scars: Ventral

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[1]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[1]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[0]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[?]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 125: Sclerites: Bivalved: Muscle scars: Ventral

0: Absent1: Present?: AmbiguousNeomorphic character.

After Bassett et al. (2001) character 6.

Alisina: Muscle scars scored based on Alisina comleyensis (Bassett et al. 2001).
Bactrotheca: “Muscle scars were not found” – Valent, Fatka, Szabad, Micka, & Marek (2012).
Halkieria evangelista: Muscle scars are known from the Type A, but not Type B, morphs of the halkieriid Oikozetetes (Jacquet, Brock, & Paterson, 2014; Paterson, Brock, & Skovsted, 2009).
Mickwitzia muralensis: Scars absent; instead, cones ornament shell’s internal surface.
Micrina: Prominent ventral muscle scars – see e.g. Holmer et al 2008, fig. 1f.
Paramicrocornus: Not preserved in conchs, though likely present (by analogy with Maxilites) (Z.-L. Zhang et al., 2018).
Tonicella: Absent (Schwabe, 2010).
Ussunia: Scars preserved on ventral valve (Nikitin & Popov, 1984).

[126] Muscle scars: Ventral: Position

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[1]Micromitra[2]Askepasma toddense[12]Coolinia pecten[12]Antigonambonites planus[?]Yuganotheca elegans[12]Tomteluva perturbata[12]Salanygolina[2]Terebratulina[-]Orthis[2]Alisina[12]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[12]Ussunia[12]Heliomedusa orienta[12]Gasconsia[-]Pelagodiscus atlanticus[-]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[12]Acanthotretella spinosa[12]Lingulosacculus[1]Clupeafumosus socialis[12]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 126: Sclerites: Bivalved: Muscle scars: Ventral: Position

1: Posterolateral and medial attachments2: Medial attachments only-: InapplicableTransformational character.

Muscles can attach to the ventral valve posterolaterally to, as well as between, the vascula lateralia (Popov 1992).

Acanthotretella spinosa: “Individual muscle scars cannot be distinguished” – Holmer & Caron 2006.
Alisina: Following reconstruction of Gorjansky & Popov (1986).
Askepasma toddense: Restricted to medial field, following the interpretation of the musculature presented by Williams et al. (2000), fig. 81.
Clupeafumosus socialis: Coded following Hadrotreta, as illustrated in Popov (1992).
Craniops: See fig. 89 in Williams et al. (2000).
Eoobolus: The ‘laterals’ of Balthasar (2009T, fig. 5) are situated almost upon the vascula lateralia; they are interpreted as sitting posterolateral to them.
Gasconsia: Musculature described in Hanken & Harper (1985), but location of mantle canals unknown.
Glyptoria: Posterolateral reflected by diductor attachments; see fig. 18.3.2 in Bassett et al. 2001.
Kutorgina chengjiangensis: Following situation in Nisusia; see fig. 18.2 in Bassett et al. (2001).
Lingulellotreta malongensis: See fig. 5 in Holmer et al. 1997E.
Micromitra: Posteriomedial muscle field (Williams et al. 1998T, text-fig. 6) treated as equivalent to posterolateral muscles.
Nisusia sulcata: Posterolateral diductors (fig. 18.2 in Bassett et al. 2001).
Novocrania: Posterior adductor muscles attach posterolaterally to ventral mantle canal (Robinson 2014).
Orthis: Not applicable: vascula lateralia not comparable to those of other taxa.
Pelagodiscus atlanticus: Inapplicable as vascular system not directly equivalent to the canonical; see. fig 6b in Balthasar (2009T).
Salanygolina: Ventral musculature not clearly constrained (Holmer et al. 2009T).
Siphonobolus priscus: Coded following general siphonotretid condition described by Popov (1992, p. 407).
Ussunia: Internal anatomy not adequately preserved to evaluate (Nikitin & Popov, 1984).

[127] Muscle scars: Adjustor

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[?]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[1]Orthis[?]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[?]Ussunia[?]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[?]Mummpikia nuda[0]Botsfordia[0]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[0]Clupeafumosus socialis[?]Micrina[?]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 127: Sclerites: Bivalved: Muscle scars: Adjustor

0: Absent1: PresentNeomorphic character.

After Bassett et al. (2001) character 7.
This character is contingent on the presence of a pedicle. Extreme caution must be used in inferring an absent state, as adjustor scars can be extremely difficult to distinguish from the adductor scars.

Alisina: Muscle scars scored based on Alisina comleyensis (Bassett et al. 2001). The presence of an adjustor is marked as not presently available, as it is not clear that a scar, if present, could be distinguished from the diminutive muscle scars present.
Askepasma toddense: Following the interpretation of the musculature presented by Williams et al. (2000), fig. 81.
Botsfordia: Not described in Popov (1992).
Clupeafumosus socialis: Not known in any acrotretid (Williams et al. 2000); not evident in Clupeafumosus (Topper et al. 2013R).
Gasconsia: No mention of an adjustor muscle in Gasconsia or Trimerellida more generally on pp. 184–185 of Williams et al. 2000, nor in discussion in Williams et al. 2007 (p. 2850). Coded as absent.
Mickwitzia muralensis: Scars absent; instead, cones ornament shell’s internal surface.
Siphonobolus priscus: Ventral musculature poorly constrained (Williams et al. 2000; Popov et al. 2009).

[128] Muscle scars: Dorsal adductors

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[12]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[?]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[2]Salanygolina[3]Terebratulina[3]Orthis[3]Alisina[?]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[3]Novocrania[3]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[3]Pelagodiscus atlanticus[-]Mickwitzia muralensis[?]Mummpikia nuda[2]Botsfordia[2]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[2]Clupeafumosus socialis[?]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 128: Sclerites: Bivalved: Muscle scars: Dorsal adductors

1: Dispersed2: Radially arranged3: Quadripartite?: Ambiguous-: InapplicableTransformational character.

After Bassett et al. (2001) character 8, and Williams et al. (1996, character 35; 2000, p. 160, character 54)

In the dorsal valve, the anterior and posterior adductor scars of articulated brachiopods form a single (quadripartite) muscle field (Williams et al. 2000, p. 201)

In contrast, the anterior and posterior scars of e.g. trimerellids have prominently separate attachment points, with anterior and posterior muscle fields clearly distinct, and coded as “dispersed”.

In e.g. kutorginates, adductor muscles are separated into left and right fields; the same is the case in lingulids, where there are more separate muscle groups and the left and right fields conspire to produce a radial arrangement; both of these configurations are scored as “radially arranged”.

Alisina: Following Williams et al. (2000) table 15 (their character 54).
Antigonambonites planus: Treatise.
Askepasma toddense: Separate left and right fields, so radially arranged – following the interpretation of the musculature presented by Williams et al. (2000), fig. 81.
Botsfordia: Following Williams et al. 1998T, appendix 2.
Clupeafumosus socialis: Following reconstruction of Hadrotreta by Williams (2000), fig. 51, which exhibits distinct left and right fields.
Coolinia pecten: “radially arranged adductor scars” – Bassett & Popov 2017, p1.
Gasconsia: Following the coding of Williams et al. (2000), table 15.

Glyptoria: Scored as “dispersed” by Williams et al. (1998T) … but then so is Kutorgina, which Bassett et al (2001) score as radial.

Williams et al. (2000) state, for superfamily Protorthida, “dorsal adductor scars probably linear”, which fits in the category of “radial” employed herein – so that’s what we follow.
Halkieria evangelista: It is unclear whether the paired muscle scars of Oikozetetes may be homologous to brachiopod adductors.
Haplophrentis carinatus: Moysiuk et al. (2017) reconstruct distinct left and right attachment scars, consistent with general situation in hyoliths (see Dzik 1980); it is unclear whether additional smaller scars were present in a radial arrangement (as in e.g. Gompholites, Marek, 1967) or whether unseen scars were dispersed, hence the partially ambiguous coding.
Heliomedusa orienta: Distinct anterior and posterior fields (Chen et al. 2007); coded as “dispersed” by Williams et al. (2000) in table 15.
Mickwitzia muralensis: Scars absent; instead, cones ornament shell’s internal surface.
Micromitra: Williams et al. (1998T) code as “dispersed”, but have a less divided scheme of character states and disagree with other sources in some codings (e.g. Bassett et al. 2001, in Kutorginates). Williams et al. (2000) do not describe Micromitra musculature and we were unable to find any reliable description of the scars, so we code as “not presently available”.
Novocrania: Craniids scored as “open, quadripartite” by Williams et al. (1996).
Pelagodiscus atlanticus: Discinids scored as “open, quadripartite” by Williams et al. (1996).
Salanygolina: “The dorsal valve of Salanygolina has a radial arrangement of adductor muscle scars and the scars of posteromedially placed internal oblique muscles, which are also characteristic of paterinates and chileates” – Holmer et al. (2009).
Siphonobolus priscus: Ventral musculature poorly constrained (Williams et al. 2000; Popov et al. 2009).
Terebratulina: Coded as “grouped, quadripartite” by Williams et al. (1996).
Ussunia: Following coding with state 0 (dispersed) in table 15 in Williams et al. 2000.

[129] Muscle scars: Adductors: Position

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[?]Micromitra[1]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[1]Salanygolina[2]Terebratulina[?]Orthis[2]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[?]Ussunia[?]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[-]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[?]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 129: Sclerites: Bivalved: Muscle scars: Adductors: Position

1: Oblique2: At high angle-: InapplicableTransformational character.

Position of adductor muscles relative to commissural plane.
After Bassett et al. (2001) character 11.

Askepasma toddense: Following the interpretation of the musculature presented by Williams et al. (2000), fig. 81.
Botsfordia: Following description of Popov (1992).
Coolinia pecten: Not reported by Williams et al. (2000), nor Bassett & Popov (2017), nor explicitly by Dewing (2001).
Eoobolus: “Eoobolus should have anterior and posterior adductors and a variety of oblique muscles which were probably arranged in criss-crossing pairs” – Balthasar 2009T.
Gasconsia: See discussion under Trimerellida in Williams et al. (2000).
Mickwitzia muralensis: Scars absent; instead, cones ornament shell’s internal surface.
Pelagodiscus atlanticus: Musculature considered essentially equivalent to Lingula by Williams et al 2000, so Lingula coding followed here.
Siphonobolus priscus: Ventral musculature poorly constrained (Williams et al. 2000; Popov et al. 2009).

[130] Muscle scars: Dermal muscles

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[?]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[?]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[0]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[?]Micrina[?]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 130: Sclerites: Bivalved: Muscle scars: Dermal muscles

0: Absent or weakly developed1: Strongly developedNeomorphic character.

Based on character 11 in Zhang et al. (2014).
Well developed dermal muscles present in the body wall of recent lingulates, which are absent in all calcareous-shelled brachiopods. These muscles are responsible for the hydraulic shell-opening mechanism, and possibly present in all organophosphatic-shelled brachiopods, with the possible exception of the paterinates (Williams et al., 2000, p. 32).

Alisina, Antigonambonites planus, Gasconsia, Glyptoria, Nisusia sulcata, Orthis, Salanygolina: According to the statement of Williams et al. (2000, p. 32) that these muscle are absent in all carbonate- shelled brachiopods.
Askepasma toddense: According to the statement of Williams et al. (2000, p. 32) that the presence of these muscles in paterinates is uncertain.
Botsfordia: Implicitly taken as present in Popov (1992), though not marked in diagrams – suggesting not strongly developed.
Clupeafumosus socialis: This character is coded based on the score of Acrotreta in Zhang et al. (2014), and statement in Williams et al. (2000, P.32).
Coolinia pecten: According to the statement of Williams et al. (2000, p. 32) that these muscle are absent in all carbonate-shelled brachiopods.
Eoobolus: Not remarked upon by Balthasar (2009T).
Kutorgina chengjiangensis: According to the statement of Williams et al. (2000, p. 32) that these muscle are absent in all carbonate- shelled brachiopods, and the coding for kutorginids in Zhang et al. (2014).
Micromitra: Williams et al. (2000, p. 32) are uncertain about the presence of these muscles in the paterinates. Zhang et al. (2014) code absence in Paterinida, but without specifying evidence; we follow their coding here.
Mummpikia nuda, Tomteluva perturbata: Though Williams et al. (2000, p. 32) state that these muscles are absent in all carbonate-shelled brachiopods, their existence cannot be discounted with certainty in this taxon, which is therefore coded not presently available.
Novocrania: Following Zhang et al. (2014), and the statement of Williams et al. (2000) that such muscles are absent in all calcite-shelled brachiopods.
Pelagodiscus atlanticus: Musculature considered essentially equivalent to Lingula by Williams et al 2000, so Lingula coding followed here.
Siphonobolus priscus: Ventral musculature poorly constrained (Williams et al. 2000; Popov et al. 2009).
Terebratulina: Williams et al. (2000, p. 32) state that these muscles are absent in all carbonate-shelled brachiopods.

[131] Muscle scars: Unpaired median (levator ani)

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[?]Micromitra[0]Askepasma toddense[0]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[?]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[?]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[?]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[0]Clupeafumosus socialis[?]Micrina[?]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 131: Sclerites: Bivalved: Muscle scars: Unpaired median (levator ani)

0: Absent1: PresentNeomorphic character.

The levator ani is a diminutive unpaired medial muscle found in certain calcitic brachiopods (Williams et al. 2000; see fig. 89, character 34 in table 13).

Alisina, Kutorgina chengjiangensis, Nisusia sulcata: Following table 13 in Williams et al. 2000.
Coolinia pecten: Not reported in Dewing (2001).
Craniops: See fig. 90 in Williams et al. 2000.
Gasconsia: Williams et al. 2000 code an unpaired medial muscle scar as present in their table 13, but give no reference for this coding, which perhaps arises from their interpretation of the taxon as a trimerellid. Hanken and Harper (1985, p. 249 and text-fig. 2) explicitly identify a pair of central muscles, so we code a levator ani as absent.
Heliomedusa orienta: Poor preservation of minor muscle scars noted by Chen et al. (2007).
Mickwitzia muralensis: Scars absent; instead, cones ornament shell’s internal surface.
Novocrania: Following table 13 in Williams et al. 2000 (for Novocrania).
Pelagodiscus atlanticus: Musculature considered essentially equivalent to Lingula by Williams et al 2000, so Lingula coding followed here.
Siphonobolus priscus: Ventral musculature poorly constrained (Williams et al. 2000; Popov et al. 2009).
Ussunia: Following table 15 in Williams et al. 2000.

[132] Muscle scars: Dorsal diductor

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[1]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[1]Ussunia[?]Heliomedusa orienta[1]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[?]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[?]Micrina[?]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 132: Sclerites: Bivalved: Muscle scars: Dorsal diductor

0: Absent1: Present?: AmbiguousNeomorphic character.

After Bassett et al. (2001) character 9.

Acanthotretella spinosa: Not observable in Acanthotretella itself, so coded as ambiguous – though it is likely based on the anticipated phylogenetic affinities of Acanthotretella that the muscles are absent.
Askepasma toddense: Possible diductor scar could instead correspond to discinoid posterior adductors (Williams et al., 1998b); coded as uncertain. Not reconstructed in the the interpretation of the musculature presented by Williams et al. (2000), fig. 81.
Clupeafumosus socialis: Not reported by Topper et al. (2013R), nor reconstructed in generic acrotretid by Williams et al. (2000).
Gasconsia: Internal oblique muscles serve as diductors.
Halkieria evangelista: It is unclear whether the paired muscle scars of Oikozetetes are homologous to brachiopod diductors.
Micromitra: Possible diductor scar could instead correspond to discinoid posterior adductors (Williams et al., 1998b); coded as uncertain.
Siphonobolus priscus: Ventral musculature poorly constrained (Williams et al. 2000; Popov et al. 2009).
Ussunia: Internal oblique muscles present (Nikitin & Popov, 1984) and taken to serve as diductors by analogy with Gasconsia.

[133] Muscle scars: Dorsal diductor: Position

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[2]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[3]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[2]Gasconsia[-]Pelagodiscus atlanticus[-]Mickwitzia muralensis[?]Mummpikia nuda[-]Botsfordia[-]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[?]Micrina[?]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 133: Sclerites: Bivalved: Muscle scars: Dorsal diductor: Position

1: Close to commissural plane2: Oblique to commissural plane3: At high angle to commissural plane-: InapplicableTransformational character.

After Bassett et al. (2001) character 10.

Siphonobolus priscus: Ventral musculature poorly constrained (Williams et al. 2000; Popov et al. 2009).

4.14 Sclerites: Dorsal valve

[134] Growth direction

Character adds 16 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[3]Tonicella[2]Mopalia[1]Polysacos vickersianum[2]Leptochiton[2]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[2]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[2]Mytilus[-]Dentalium[2]Pelagiella[1]Siphogonuchites multa[1]Acaenoplax hayae[2]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[1]Orthrozanclus[2]Micromitra[2]Askepasma toddense[3]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[2]Tomteluva perturbata[2]Salanygolina[2]Terebratulina[2]Orthis[2]Alisina[2]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[2]Ussunia[1]Heliomedusa orienta[2]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[2]Mummpikia nuda[2]Botsfordia[2]Eoobolus[2]Lingulellotreta malongensis[3]Lingula[2]Siphonobolus priscus[2]Acanthotretella spinosa[2]Lingulosacculus[13]Clupeafumosus socialis[2]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 134: Sclerites: Dorsal valve: Growth direction

1: Holoperipheral2: Mixoperipheral3: Hemiperipheral?: Ambiguous-: InapplicableTransformational character.

See Fig. 284 in Williams et al (1997).
The growth direction dictates the attitude of the cardinal area relative to the hinge, which does not therefore represent an independent character.
Crudely put, if, viewed from a dorsal position, the umbo falls within the outer margin of the shell, growth is holoperipheral; if it falls outside the margin, it is mixoperipheral; if it falls exactly on the margin, it is hemiperipheral.

Acaenoplax hayae: Holoperipheral (???).
Calvapilosa kroegeri: Umbo in centre of valve (Vinther et al., 2017).
Clupeafumosus socialis: Appears hemiperipheral in fig. 3 in Topper et al (2013R), though bordering on holoperipheral, so scored as ambiguous.
Craniops: “Both valves with growth holoperipheral” – Williams et al. 2000, p. 164.
Glaphurochiton carbonarius: Seemingly mixoperipheral (Hoare & Mapes, 1986).

Heliomedusa orienta: “holoperipheral growth in dorsal valve” – Williams et al. 2007.

Zhang et al. (2009) conclude that Chen et al. (2007) misidentify the dorsal valve as the ventral valve.
Kulindroplax perissokomos: “All valves are mixoperipheral” (Sutton et al., 2012).
Micrina: See Holmer et al. (2008).
Neopilina: Mixoperipheral, though only just (Menzies & Layton, 1962).
Orthrozanclus: The umbo is situated in the anterior half of the shell, but within the margin (Conway Morris & Caron, 2007).
Paterimitra: S2 and L sclerites are clearly holoperipheral. See Larsson et al. 2014, fig. 2.
Phthipodochiton thraivensis: All valves mixoperipheral (Sutton & Sigwart, 2012).
Siphogonuchites multa: Bengtson (1992).

Tonicella: For the purposes of this analysis, we must treat polyplacophoran and brachiopod valves as potentially homologous.

In brachiopods, the dorsal valve bears the lophophore, which arises from the anterior lobe of the larva (Altenburger et al., 2013) – indicating that the dorsal shell field is associated with the anterior lobe.

In polyplacophorans, the head valve arises from a shell field on the anterior (pre-prototroch) lobe of the larva (Wanninger & Haszprunar, 2002a), which we therefore treat as homologous with the brachiopod dorsal valve.

In support of this hypothesis, we note that the posterior (but not anterior) valves of chitons bear apophyses (Connors et al., 2012; Schwabe, 2010), which are most prominent in the ventral (but not dorsal) valves of brachiopods (Williams et al 1997, fig. 322), and which occur in the morph A shell of Oikozetetes, which is interpreted as the posterior valve of a halkieriid (Paterson et al., 2009).

As the single posterior shell field of polyplacophorans subdivides to give rise to the six intermediate valves plus the tail valve (Wanninger & Haszprunar, 2002a), we prefer to consider the intermediate valves as representing “subdivisions” of a single valve rather than additional valves added to the body plan.

Growth is hemiperipheral in the anterior valve of polyplacophorans and holoperipheral in the posterior valves (Connors et al., 2012; Schwabe, 2010).
Ussunia: Growth “mixoperipheral in both valves” in trimerellids (Popov, Holmer, & Gorjansky, 1997; Williams et al., 2000).

[135] Aspect

Character adds 7 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[3]Tonicella[3]Mopalia[3]Polysacos vickersianum[3]Leptochiton[23]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[3]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[?]Dentalium[1]Pelagiella[1]Siphogonuchites multa[2]Acaenoplax hayae[3]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[2]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[3]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[3]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 135: Sclerites: Dorsal valve: Aspect

1: Elongate: longer than wide2: Equant: length ~ width3: Transverse: wider than long?: Ambiguous-: InapplicableTransformational character.

Length:width ratio of the primary valve. Coded ambiguous in marginal cases: for example, a length:width ratio of 1.02:1 might be coded ambiguous(elongate, equant).

Acaenoplax hayae: Equant (???).
Calvapilosa kroegeri: Slightly longer than wide (Vinther et al., 2017).
Glaphurochiton carbonarius: Described as “semi-circular to elongate”, but some figured material is almost wider than long (Hoare & Mapes, 1986); coded as equant to transverse.
Halkieria evangelista: Approximately equant (???).
Kulindroplax perissokomos: Wider than long (Sutton et al., 2012).
Mopalia: (???).
Novocrania: Wider than long (???).
Pelagiella: Nützel et al. (2006).
Pelagodiscus atlanticus: Approximately equant (???).
Siphogonuchites multa: Bengtson (1992).

[136] Posterior surface: Differentiated

Character adds 9 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[0]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[1]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[1]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 136: Sclerites: Dorsal valve: Posterior surface: Differentiated

0: Posterior face of dorsal valve not differentiated1: Posterior face of dorsal valve forms distinct cardinal area or pseudointerarea?: AmbiguousNeomorphic character.

In shells that grow by mixoperipheral growth, the triangular area subtended between each apex and the posterior ends of the lateral margins is termed the cardinal area. In shells with holoperipheral growth, a flattened surface on the posterior margin of the valve is termed a pseudointerarea (paraphrasing Williams et al. 1997).

In order for this character to be independent of a shell’s growth direction, we do not distinguish between a “cardinal area”, “interarea” or “pseudointerarea”.

Acaenoplax hayae: Not differentiated; essentially round (???).
Acanthotretella spinosa: Pseudointerarea present, following Siphonotretidae coding in Williams et al. (2000), table 6.
Alisina, Antigonambonites planus, Coolinia pecten, Glyptoria, Kutorgina chengjiangensis, Orthis, Salanygolina, Tomteluva perturbata: Cardinal area (interarea) present.
Askepasma toddense: Well-defined pseudointerarea (Williams et al. 2000, p153).
Botsfordia: “dorsal pseudointerarea vestigial, divided by median groove” – Williams et al. 2000.
Calvapilosa kroegeri: Slight concavity of posterior surface (Vinther et al., 2017).
Clupeafumosus socialis: Pseudointerarea present; figured by Topper et al. (2013R), fig. 3j.
Craniops: “Only some craniopsids (Lingulapholis, Pseudopholidops [not Craniops]) have well-developed pseudointerareas.” – Williams et al. 2000.
Gasconsia: Absent: the dorsal (branchial) pseudointerarea of G. schucherti is “reduced or obsolete”; that of G. worsleyi “short, virtually obsolete” (Hanken & Harper 1985).
Haplophrentis carinatus: A very short pseudointerarea appears to be present (Moysiuk et al. 2017).
Heliomedusa orienta: Pseudointerea in ventral valve, but not dorsal valve (Williams et al. 2000, 2007).
Leptochiton: (???).
Lingula, Lingulellotreta malongensis: Pseudointerarea present, following Williams et al. (2000), table 6.
Lingulosacculus: Unclear from fossil material.
Longtancunella chengjiangensis: Zhang et al. (2011T) note that “all evidence of a pseudointerarea is lacking”, but the two-dimensional preservation style of Chengjiang material makes details of dorsal valve difficult to distinguish, and the possibility of a diminutive pseudointerarea cannot be excluded with total confidence.
Mickwitzia muralensis: Shell flat.
Micrina: = Sellate sclerite duplicature (Holmer et al. 2008).
Micromitra: “Dorsal pseudointerarea usually well defined, low, anacline to catacline” – Williams et al. 2000.
Mummpikia nuda: “Information on the dorsal interarea is inconclusive […] no obvious
interarea is recognisable; whether or not this is the primary state or a taphonomic artefact is difficult to assess” – Balthasar 2008, p. 276.
Nisusia sulcata: Cardinal area (interarea) present – with reference to Holmer et al. (2018E).
Novocrania, Paterimitra, Pedunculotheca diania: Pseudointerarea.
Pelagodiscus atlanticus: Absent, following entry for Discinidae in Williams et al. (2000), table 6.
Phthipodochiton thraivensis: Slightly concave (Sutton & Sigwart, 2012).
Siphogonuchites multa: Bengtson (1992).
Siphonobolus priscus: “Dorsal pseudointerarea weakly anacline, undivided, elevated above the valve floor” – Popov et al. 2009.
Terebratulina: Interarea present.
Tonicella: V-shaped notch in anterior valve (Schwabe, 2010).
Ussunia: Following table 15 in Williams et al. 2000.
Yuganotheca elegans: A differentiated region is not obvious in fossil material or its reconstruction (Zhang et al. 2014), but the two-dimensional preservation style of Chengjiang material makes details of dorsal valve difficult to distinguish, and the possibility of a diminutive pseudointerarea cannot be excluded with confidence.

[137] Differentiated posterior surface: Morphology

Character adds 12 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[2]Mopalia[0]Polysacos vickersianum[2]Leptochiton[2]Glaphurochiton carbonarius[2]Calvapilosa kroegeri[2]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[2]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[1]Haplophrentis carinatus[0]Cupitheca holocyclata[01]Bactrotheca[01]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[1]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[0]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[?]Mummpikia nuda[0]Botsfordia[1]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[01]Siphonobolus priscus[0]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[0]Micrina[1]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 137: Sclerites: Dorsal valve: Differentiated posterior surface: Morphology

0: Convex lateral profile1: Planar lateral profile2: Concave lateral profile?: AmbiguousNeomorphic character.

It is possible for a cardinal area or pseudointerarea to be distinct from the anterior part of the shell, yet to remain curved in lateral profile.

Taking an undifferentiated posterior margin as primitive, the primitive condition is curved – flattening of the posterior margin represents an additional modification that can only occur once the posterior margin is differentiated.

Bactrotheca: The short aspect of the cardinal interarea (Valent et al., 2012) makes it difficult to evaluate whether it is planar or curved.
Botsfordia: “Curved pseudointerarea” – Skovsted et al. 2017.
Calvapilosa kroegeri: Slight concavity of posterior surface (Vinther et al., 2017).
Clupeafumosus socialis: Truncated but essentially planar surface; see e.g. p196 of Topper et al. 2013R.
Cupitheca holocyclata: Curved (H.-J. Sun et al., 2018).
Eoobolus: Essentially planar; see Balthasar (2009T), fig. 4a.
Pelagodiscus atlanticus, Gasconsia, Heliomedusa orienta, Mickwitzia muralensis, Ussunia: Posterior surface cannot be flat if it is not differentiated.
Glaphurochiton carbonarius: Irregular but overall concave (Hoare & Mapes, 1986).
Micromitra: Essentially straight; see fig. 3.7 in Ushatinskaya 2016P.
Orthrozanclus: Posterior face appears close to planar (Conway Morris & Caron, 2007; Zhao et al., 2017).
Paramicrocornus: Z.-L. Zhang et al. (2018).
Pedunculotheca diania: Difficult to evaluate based on present material, given low nature of valve and compressed preservation.
Siphonobolus priscus: The short interarea appears planar (see for example Popov et a. 2009 fig. 6A), but its short length makes it difficult to establish whether slight curvature is present.
Tonicella: Essentially planar, though open in aspect (following Chiton in Schwabe, 2010).

[138] Posterior surface: Medial groove

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[1]Conocardium elongatum[0]Haliotis[1]Pojetaia runnegari[1]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[?]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[?]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 138: Sclerites: Dorsal valve: Posterior surface: Medial groove

0: Absent1: Present?: AmbiguousNeomorphic character.

Following character 29 in Williams et al. (2000), table 9 (which relates to pseudointerarea).

Acanthotretella spinosa: The dorsal pseudointerarea is poorly preserved, but appears to have a median groove (Holmer & Caron, 2006).
Botsfordia: “dorsal pseudointerarea vestigial, divided by median groove” – Williams et al. 2000.
Clupeafumosus socialis: Present; figured by Topper et al. (2013R), fig. 3j.
Conocardium elongatum: Arguably the gap between the valves represents a medial groove.
Eoobolus: Prominent medial groove (Balthasar 2009T).
Heliomedusa orienta: “A posteriorly protruding dorsal pseudointerarea with no median groove and no flexure lines” – Chen et al. 2007.
Lingulellotreta malongensis: Dorsal pseudointerarea with wide, concave median groove and short propareas" – Williams et al 2000.
Pojetaia runnegari, Mytilus: The gap between the valves could arguably be described as a medial groove.
Paramicrocornus: Z.-L. Zhang et al. (2018).
Siphonobolus priscus: The dorsal pseudointerarea of S. priscus is undivided (Popov et al. 2009), but in other species it is divided by a “wide, poorly defined median groove” (Williams et al. 2000). Coded, therefore, as polymorphic.

[139] Posterior surface: Notothyrium

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[0]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[?]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 139: Sclerites: Dorsal valve: Posterior surface: Notothyrium

0: Absent1: PresentNeomorphic character.

A notothyrium is an opening in an interarea that accommodates the pedicle, and may be filled with plates.

Botsfordia: Following Williams et al. 1998T, appendix 2.
Longtancunella chengjiangensis: No evidence or report of an opening at the hinge line in fossil material in Zhang et al. 2007A or Zhang et al. 2011T.
Paramicrocornus: Z.-L. Zhang et al. (2018).
Tonicella: The deep V-shaped notch (Schwabe, 2010, fig. 8) is positionally equivalent to the brachiopod notothyrium.

[140] Posterior surface: Notothyrium: Shape

Character adds 2 to tree score (1 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[-]Micromitra[-]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[-]Yuganotheca elegans[2]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[2]Orthis[1]Alisina[-]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[-]Pelagodiscus atlanticus[-]Mickwitzia muralensis[?]Mummpikia nuda[-]Botsfordia[-]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[-]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 140: Sclerites: Dorsal valve: Posterior surface: Notothyrium: Shape

1: Parallel-sided cleft2: Triangular-: InapplicableTransformational character.

A notothyrium is an opening in an interarea that accommodates the pedicle, and may be filled with plates.

A simplification of character 5 in Bassett et al. 2001.

[141] Posterior surface: Notothyrium: Chilidial plates

Character adds 2 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[-]Micromitra[-]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[-]Yuganotheca elegans[2]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[1]Orthis[1]Alisina[-]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[-]Pelagodiscus atlanticus[-]Mickwitzia muralensis[?]Mummpikia nuda[-]Botsfordia[-]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[-]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 141: Sclerites: Dorsal valve: Posterior surface: Notothyrium: Chilidial plates

1: Open2: Covered by chilidial plates?: Ambiguous-: InapplicableTransformational character.

A notothyrium may be open or covered by a chilidium or two chilidial plates.
No included taxa exhibit more than one chilidial plate.
Transformational as it is not self-evident whether the ancestral taxon had an open or closed notothyrium.

[142] Notothyrial platform

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[?]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 142: Sclerites: Dorsal valve: Notothyrial platform

0: Absent1: Present?: AmbiguousNeomorphic character.

After Bassett et al. (2001) character 12.
The presence or absence of a notothyrial platform, which often serves as an attachment point for the diductors in a similar fashion to the cardinal processes, is independent of the presence of a notothyrium.

Alisina, Glyptoria: Bassett et al. (2001) score as present in Table 18.1.
Askepasma toddense: Raised notothyrial platform (Williams et al., 1998b).
Coolinia pecten: Referred to as the “posterior platform” in Dewing (2001).
Kutorgina chengjiangensis: “Dorsal diductor scars impressed on floor of notothyrial cavity”: Williams et al. 2000, regarding Kutorginata.
Bassett et al. (2001) score as absent in Table 18.1.
Micromitra: A low notothyrial plate (Williams et al., 1998b) conceivably correspond to the raised notothyrial platform of Askepasma; coded ambiguous accordingly.
Nisusia sulcata: Bassett et al. (2001) score as absent in Table 18.1.
“Dorsal diductor scars impressed on floor of notothyrial cavity”: Williams et al. 2000, regarding Kutorginata.
Ussunia: “Visceral platforms absent in both valves” – Williams et al. 2000, p. 192.

[143] Medial septum

Character adds 6 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[1]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[1]Novocrania[0]Craniops[0]Ussunia[1]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 143: Sclerites: Dorsal valve: Medial septum

0: Absent1: PresentNeomorphic character.

The dorsal valve of many taxa is exhibits a septum or process (or myophragm) along the medial line. See character 25 in Benedetto (2009).

Acanthotretella spinosa: Not described by Holmer & Caron (2006), but an unannotated linear feature corresponds to the position of a median septum. Without detailed study of the specimen, we opt to score this as ambiguous.
Antigonambonites planus: Weakly developed septum evident in internal cast: Williams et al. 2000, fig. 508.2e.
Botsfordia: “dorsal interior with narrow anterior projection extending to midvalve, bisected by median ridge” – Williams et al. 2000.
Clupeafumosus socialis: Prominent process evident (Topper et al., 2013R).
Eoobolus: A “median projection” is present (fig. 4g in Balthasar 2009T).
Glyptoria: Neither evident nor reported in Williams et al. (2000).

Heliomedusa orienta: Reported on ‘ventral’ valve by Chen et al. (2007); we consider their ‘ventral’ valve to be the dorsal valve.

The structure is unambiguously figured (e.g. fig. 5.1 in Chen et al. 2007), contra its coding as absent in Williams et al. 2000 and its lack of mention in Williams et al. 2007 or Zhang et al. 2009.
Kutorgina chengjiangensis: Absent – fig. 129.1f in Williams et al. (2000).
Lingulellotreta malongensis: Very weakly developed but seemingly present between muscle scars in Lingulellotreta, more prominent in Aboriginella (also Lingulellotretidae) (Williams et al. 2000, fig. 34).
Lingulosacculus: It is not possible to determine, based on the material presented in Balthasar & Butterfield (2009E), whether the anterior projection of the visceral area in the dorsal valve corresponds to a medial septum in the underlying shell.
Mummpikia nuda: See pl. 2 panel 6 in Balthasar (2008).
Nisusia sulcata: Fig. 125 in Williams et al. (2000).
Novocrania: Median process evident: Williams et al. (2000) fig. 100.2a, d.
Orthis: Short medial process (“low median ridge”, p. 724) present in dorsal valve; see Fig. 523.3b in Williams et al. (2000).
Paramicrocornus: Z.-L. Zhang et al. (2018).
Siphonobolus priscus: “Dorsal interior […] bisected by a short median ridge.” – Popov et al. 2009.
Ussunia: Following char 42 in table 15 in Williams et al. 2000.

[144] Cardinal shield

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[0]Bactrotheca[1]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 144: Sclerites: Dorsal valve: Cardinal shield

0: Absent1: Present?: AmbiguousNeomorphic character.

The hyolithid operculum is divided into a cardinal and conical shield (Z.-L. Zhang et al., 2018), separated by furrows corresponding to the position of the helens. See Marek (1976) (fig. 2) or Martí Mus & Bergström (2005) (fig. 1) for schematic.

With no obvious sites for muscle attachment, the shields are unlikely to be homologous to the notothyrial platform.

Bactrotheca: No differentiation between the cardinal and conical shields.
Cupitheca holocyclata: Narrow cardinal shield (Skovsted et al., 2016).
Paramicrocornus: Z.-L. Zhang et al. (2018).

[145] Cardinal processes

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[?]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[0]Yuganotheca elegans[1]Tomteluva perturbata[0]Salanygolina[1]Terebratulina[1]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 145: Sclerites: Dorsal valve: Cardinal processes

0: Absent1: Present?: AmbiguousNeomorphic character.

After Bassett et al. (2001) character 13. See Martí Mus & Bergström (2005) for an illustration.
Cardinal processes are unlikely to be homologous with the notothyrial platform, even if their function is similar.

Bactrotheca: “Narrow broadly diverging cardinal processes with subparallel edges” – Valent et al. (2012).
Clupeafumosus socialis: Not reported by Topper et al. (2013R).
Cupitheca holocyclata: Well-defined (Skovsted et al., 2016).
Longtancunella chengjiangensis: Not evident, and ought arguably to be discernable if present given the quality of preservation.
Paramicrocornus: Present (Z.-L. Zhang et al., 2018).

[146] Cardinal teeth

Character adds 0 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 146: Sclerites: Dorsal valve: Cardinal teeth

0: AbsentNeomorphic character.

Radially arranged teeth, separated by furrows, adorn the cardinal margin of the operculum of certain hyolithids (Marek, 1963). The absence of corresponding tooth sockets indicates that they do not serve to articulate the valves; Marek (1967) does not consider the teeth to be homologous with brachiopod cardinal teeth.

[147] Clavicles

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 147: Sclerites: Dorsal valve: Clavicles

0: Absent1: PresentNeomorphic character.

Prominent symmetrical ridges on the inner surface of the hyolith operculum.

Cupitheca holocyclata: “No traces of clavicles” (Skovsted et al., 2016).
Paramicrocornus: Z.-L. Zhang et al. (2018).

[148] Clavicles: Type of clavicles

Character adds 0 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[-]Micromitra[-]Askepasma toddense[-]Coolinia pecten[-]Antigonambonites planus[-]Yuganotheca elegans[-]Tomteluva perturbata[-]Salanygolina[-]Terebratulina[-]Orthis[-]Alisina[-]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[-]Pelagodiscus atlanticus[-]Mickwitzia muralensis[-]Mummpikia nuda[-]Botsfordia[-]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[-]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 148: Sclerites: Dorsal valve: Clavicles: Type of clavicles

1: Monoclavicle-: AmbiguousTransformational character.

Usually the operculum of hyoliths has one pair of clavicles, but in some taxa of hyolithida there are more than one pair of clavicles, which can be divided into six types (Marek, 1967). The included taxa either exhibit a single pair of monoclavicles, or three pairs of clavicles.

Paramicrocornus: Z.-L. Zhang et al. (2018).

4.15 Sclerites: Ventral valve

[149] Growth direction

Character adds 6 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[-]Polysacos vickersianum[1]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[2]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[2]Salanygolina[2]Terebratulina[2]Orthis[2]Alisina[2]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[2]Ussunia[2]Heliomedusa orienta[2]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[2]Mummpikia nuda[2]Botsfordia[2]Eoobolus[2]Lingulellotreta malongensis[3]Lingula[12]Siphonobolus priscus[2]Acanthotretella spinosa[2]Lingulosacculus[12]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 149: Sclerites: Ventral valve: Growth direction

1: Holoperipheral2: Mixoperipheral3: Hemiperipheral?: Ambiguous-: InapplicableTransformational character.

See Fig. 284 in Williams et al. (1997) for depiction of terms.

The growth direction dictates the attitude of the cardinal area relative to the hinge, which does not therefore represent an independent character.
Crudely put, if, viewed from a dorsal position, the umbo falls within the outer margin of the shell, growth is holoperipheral; if it falls outside the margin, it is mixoperipheral; if it falls exactly on the margin, it is hemiperipheral.

Clupeafumosus socialis: Inferred from Topper et al. (2013R).
Craniops: “Both valves with growth holoperipheral” – Williams et al. 2000, p. 164.
Heliomedusa orienta: Williams et al. (2000, 2007) reconstruct mixoperipheral growth in the ventral valve (though Chen et al. (2007) reconstruct the valves the other way round, i.e. it is the ventral valve that grows holoperipherally, and the dorsal mixoperipherally).
Paterimitra: The apical flange notwithstanding, the umbo of the S1 sclerite is posterior of the hinge line and the posterior edge of the lateral plate – see Larsson et al. 2014, fig. 2a, c.
Siphonobolus priscus: Initially holoperipheral (Popov et al. 2009, p. 159), then on the brink of being mixoperipheral in adulthood, so coded as polymorphic.
Tonicella: Growth is hemiperipheral in the anterior valve of polyplacophorans and holoperipheral in the posterior valves (Connors et al., 2012; Schwabe, 2010).
Ussunia: Growth “mixoperipheral in both valves” in trimerellids (Popov et al., 1997; Williams et al., 2000).

[150] Relative size

Character adds 11 to tree score (10 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[12]Tonicella[1]Mopalia[-]Polysacos vickersianum[23]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[1]Micromitra[1]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[12]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[2]Terebratulina[2]Orthis[2]Alisina[12]Longtancunella chengjiangensis[2]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[3]Novocrania[2]Craniops[2]Ussunia[1]Heliomedusa orienta[12]Gasconsia[3]Pelagodiscus atlanticus[1]Mickwitzia muralensis[2]Mummpikia nuda[1]Botsfordia[23]Eoobolus[1]Lingulellotreta malongensis[2]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 150: Sclerites: Ventral valve: Relative size

1: Ventral valve markedly larger than dorsal valve (ventribiconvex)2: Equivalve (subequally biconvex)3: Dorsal valve markedly larger than ventral valve (dorsibiconvex)?: Ambiguous-: InapplicableTransformational character.

In many brachiopods, the valves are closely similar in size; in others, the ventral valve is markedly larger than the dorsal, on account of being more convex. Marginal cases are treated as ambiguous for the relevant states.

Antigonambonites planus: Broadly equivalve – see Williams et al. (2000) fig. 508.2c.
Botsfordia: After table 8 in Williams et al. (2000).
Craniops: “Shell subequally biconvex” – Williams et al. 2000.
Eoobolus: “Eoobolus is biconvex”, but in his amended diagnosis, Balthasar (2009T) described it as “shell inequivalved, dorsibiconvex”.
Gasconsia: Equivalve as juveniles, becoming “convexiplane” (Williams et al. 2000, p. 187) as adults (Hanken & Harper, 1985).
Heliomedusa orienta: Ventral valve larger than the dorsal valve (Zhang et al. 2009, p. 659).
Kutorgina chengjiangensis: Ventral valve larger (see Williams et al. 2000, fig. 125.).
Leptochiton: Dorsal valve slightly larger (???).
Longtancunella chengjiangensis, Yuganotheca elegans: The ventral valve is somewhat, but not markedly, larger than the dorsal; as such, this character is coded ambiguous for equivalve/ventral valve larger.
Mummpikia nuda: Aside from hinge, valves similar in convexity and size (Balthasar 2008).
Nisusia sulcata: Ventral valve larger (see Williams et al. 2000, fig. 126.).
Siphonobolus priscus: Ventribiconvex (Popov et al. 2009).
Tonicella: Coded as ambiguous for equivalve/ventral valve larger: the posterior embryonic shell field, treated herein as equivalent to the ventral valve,.
Ussunia: Subequally biconvex (Williams et al. 2000, p. 192).

[151] Posterior surface: Differentiated

Character adds 7 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[0]Polysacos vickersianum[1]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[?]Pedunculotheca diania[1]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[1]Heliomedusa orienta[1]Gasconsia[0]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 151: Sclerites: Ventral valve: Posterior surface: Differentiated

0: Posterior surface of shell not differentiated1: Posterior surface of shell forms distinct cardinal area or pseudointerareaNeomorphic character.

In shells that grow by mixoperipheral growth, the triangular area subtended between each apex and the posterior ends of the lateral margins is termed the cardinal area. In shells with holoperipheral growth, a flattened surface on the posterior margin of the valve is termed a pseudointerarea (paraphrasing Williams et al. 1997).

In order for this character to be independent of a shell’s growth direction, we do not distinguish between a “cardinal area”, “interarea” or “pseudointerarea”.

Alisina, Antigonambonites planus, Coolinia pecten, Glyptoria, Kutorgina chengjiangensis, Nisusia sulcata, Orthis, Salanygolina, Tomteluva perturbata: Interarea present.
Bactrotheca: No clear delineation of posterior (functionally dorsal) surface.
Clupeafumosus socialis: Described by Topper et al. (2013R).
Cupitheca holocyclata: No evidence of differentiation; circular cross-section (Vendrasco, Checa, & Porter, 2017).
Gasconsia: The region corresponding to the ventral (pseudo)interarea is described as a “trimerellid ventral cardinal area” by Williams et al. (2000, p.162), who code both an interarea and a pseudointerarea as absent in trimerellids.
Heliomedusa orienta: Zhang et al. (2009) report a moderate to somewhat developed ventral pseudointerarea, confirmed by Williams et al (2007).
Lingulosacculus: The conical valve is interpreted as the ventral valve with an extended pseudointerarea.
Longtancunella chengjiangensis: Though “all evidence of a pseudointerarea is lacking” – Zhang et al. 2011T – the region of the shell between the strophic hinge line and the colleplax (fig. 2 in Zhang et al. 2011T) is distinct from the rest of the shell; the ends of the strophic hinge line are marked by prominent nicks in the shell margin. Longtancunella is therefore coded as having a differentiated posterior surface.
Mickwitzia muralensis: Termed an interarea by Balthasar (2004).
Mopalia: (???).
Mummpikia nuda: Balthasar (2008) interprets a pseudointerarea as being present – e.g. p273, “Of particular interest is the vault that bridges the most anterior portion of the ventral pseudointerarea and raises it above the visceral platform.”; “This pattern is reversed in the ventral valves of M. nuda, where the anterior projection of the pedicle groove is raised above the valve floor whereas the lateral parts of pseudointerarea are not”.
Paterimitra: Triangular notch and subapical flange.
Pedunculotheca diania: Lateral lines suggest differentiation of posterior surface, but difficult to discern a change in morphology of this region. Coded ambiguous.
Siphonobolus priscus: “Ventral pseudointerarea, low, undivided, poorly defined” – Williams et al. 2000.
Terebratulina: Interarea.
Tonicella: Following the proposed homology model between the posterior valve of polyplacophorans and the ventral valve of brachiopods, the “posterior” surface of the polyplacophoran valve is taken to be the surface that would articulate with the anterior valve, which is anatomically anterior on the living organism.
Ussunia: Following char 17 in table 15 of Williams et al. 2000.

[152] Posterior margin growth direction

Character adds 3 to tree score (2 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[-]Polysacos vickersianum[2]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[2]Paramicrocornus[2]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[2]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[2]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[2]Yuganotheca elegans[2]Tomteluva perturbata[2]Salanygolina[2]Terebratulina[2]Orthis[2]Alisina[2]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[2]Heliomedusa orienta[2]Gasconsia[-]Pelagodiscus atlanticus[2]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[2]Lingula[2]Siphonobolus priscus[2]Acanthotretella spinosa[?]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 152: Sclerites: Ventral valve: Posterior margin growth direction

1: Inward-growing2: Outward-growing-: InapplicableTransformational character.

Balthasar (2008) notes an inward-growing posterior margin of the pseudointerarea as potentially linking Mummpikia with the linguliform brachiopods.

Coded as inapplicable in taxa without a differentiated posterior margin: the posterior margin can only grow inwards if it is differentiated from the anterior margin; else the entire shell would grow in on itself.

Botsfordia: Inward-growing; see Skovsted & Holmer (2005), pl. 4.
Clupeafumosus socialis: See Topper et al. (2013R).
Eoobolus: See for example Skovsted & Holmer (2005), pl. 3.
Lingulellotreta malongensis: Transverse cross section of ventral pseudointerarea concave.
Mopalia: (???).
Mummpikia nuda: Balthasar (2008) interprets an inward-growing posterior margin of the pseudointerarea – e.g. p273, “Of particular interest is the vault that bridges the most anterior portion of the ventral pseudointerarea and raises it above the visceral platform […] An inward-growing posterior margin is otherwise known only from the pseudointerareas of linguliform brachiopods”.

[153] Posterior surface: Planar

Character adds 8 to tree score (7 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[-]Polysacos vickersianum[0]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[1]Pedunculotheca diania[1]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[1]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[1]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[01]Siphonobolus priscus[1]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 153: Sclerites: Ventral valve: Posterior surface: Planar

0: Curved lateral profile1: Planar lateral profile?: Ambiguous-: InapplicableNeomorphic character.

It is possible for a cardinal area or pseudointerarea to be distinct from the anterior part of the shell, yet to remain curved in lateral profile.

Taking an undifferentiated posterior margin as primitive, the primitive condition is curved – flattening of the posterior margin represents an additional modification that can only occur once the posterior margin is differentiated.

A flat and triangular interarea links Mummpikia with the Obolellidae (Balthasar 2008) – but all included taxa have triangular interareas, so this is not listed as a separate character.

Acanthotretella spinosa: ventral pseudointerareas are most similar to those found within the Order Siphonotretida.
Botsfordia: See Skovsted & Holmer (2005), pl. 3, fig. 14.
Clupeafumosus socialis: “Ventral pseudointerarea is gently procline and is flat in lateral profile”. —
(Topper et al. 2013R).
Eoobolus: Some curvature retained.
Haplophrentis carinatus: Dorsal surface essentially linear (Moysiuk et al., 2017, fig ed6a, ed7a).
Lingulellotreta malongensis: Transverse cross section of ventral pseudointerarea concave.
Longtancunella chengjiangensis: Flattened, reflecting the strophic hinge line.
Micromitra: Essentially planar; see fig. 6 in Ushatinskaya 2016P.
Mopalia: (???).
Paramicrocornus: Posterior (functionally dorsal) surface is linear in adults (though slightly curving in juveniles) (Z.-L. Zhang et al., 2018).
Pedunculotheca diania: Essentially linear.
Siphonobolus priscus: ‘Almost’ planar – see Popov et al. (2009, fig. 4). Coded as ambiguous.
Tonicella: (Schwabe, 2010).

Oh dear! You included the inapplicable token in a neomorphic character!
That’s unlikely to be a good idea: see Brazeau, Smith & Guillerme (2018, Systematic Biology).
Unless you are very sure that you understand the consequences, you should mark the character as Transformational by setting State 0 to [Transformational character], or re-code:

  • Acaenoplax_hayae
  • Kulindroplax_perissokomos
  • Phthipodochiton_thraivensis
  • Calvapilosa_kroegeri
  • Glaphurochiton_carbonarius
  • Polysacos_vickersianum

[154] Ligula

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 154: Sclerites: Ventral valve: Ligula

0: Absent1: PresentNeomorphic character.

The aperture of many hyolithid hyoliths is characterised by a ligula, a tongue-like protruding shelf on the functionally ventral surface of conical shell (Martí Mus & Bergström, 2005). This can be recognized by an acute angle in the lateral profile of the commissure (see second figure on p. 91 of Marek, 1966). No brachiopods display an equivalent feature.

Paramicrocornus: “Very short” semicircular ligula (Z.-L. Zhang et al., 2018).

[155] Posterior surface: Extent

Character adds 6 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[2]Paramicrocornus[2]Haplophrentis carinatus[-]Cupitheca holocyclata[2]Bactrotheca[2]Pedunculotheca diania[1]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[12]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[12]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[12]Nisusia sulcata[12]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[12]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[2]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[2]Acanthotretella spinosa[2]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 155: Sclerites: Ventral valve: Posterior surface: Extent

1: Low: Wider than deep2: High: Deeper than wide?: Ambiguous-: InapplicableTransformational character.

Distinguishes taxa whose ventral valve is essentially flat from those that are essentially conical.

Antigonambonites planus: Though scored High in data matrix of Benedetto (2009), this taxon (see Williams et al. 2000, fig. 508) does not express the deeply conical ventral valve that this character is intended to reflect. It is charitably coded as ambiguous.
Clupeafumosus socialis: Entire valve length – see schematic in Williams et al. (1997), fig. 286.
Coolinia pecten: See fig. 485 in Williams et al. 2000.
Gasconsia: “ventral cardinal interarea low, apsacline, with narrow, poorly defined homeodeltidium” – Williams et al. 2000, p. 186.
Kutorgina chengjiangensis: This taxon (see Williams et al. 2000, fig. 129; Popov 1992, fig. 1) comes close to expressing the deeply conical ventral valve that this character is intended to reflect, though this is not always so pronounced (e.g. Williams et al. 2000, fig. 125). It is therefore coded as ambiguous.
Mickwitzia muralensis: Often not prominently high (Skovsted & Holmer, 2003; Balthasar, 2004), though in some cases (e.g. Butler et al. 2015) the ventral valve approaches the conical shape that this character is intended to capture. Coded as polymorphic.
Nisusia sulcata: Scored as high in data matrix of Benedetto (2009), and depicted as such in Williams et al. (2000, fig. 125) and Popov (1992, fig. 1); but not high in all specimens (e.g. Williams et al. 2000, fig. 126). It is therefore coded as polymorphic.
Novocrania: Low cone.
Orthis: Scored ‘Low’ for Eoorthis by Benedetto (2009); assumed same in Orthis.
Salanygolina: Whereas Williams et al. (2000, p. 156) describe the ventral pseudointerarea as high, the shell lacks the deeply conical aspect that this character is intended to capture; we thus code the taxon as ambiguous.

[156] Posterior surface: Delthyrium

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[0]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 156: Sclerites: Ventral valve: Posterior surface: Delthyrium

0: Absent1: PresentNeomorphic character.

A delthyrium is an opening in an interarea or pseudointerarea that accommodates the pedicle, and may be filled with plates.

The homology of the pedicle in the pseudointerarea of obolellids and botsfordiids with the umbonal pedicle foramen of acrotretids was proposed by Popov (1992), and seemingly corroborated by observations of Ushatinskaya & Korovnikov (2016R), who note that the propareas of the Botsfordia ventral valve sometimes merge to form an elongate teardrop-shaped pedicle foramen.

Acanthotretella spinosa: Origin modelled on Siphonobolus.
Askepasma toddense: Homeodeltidium absent (Williams et al. 2000, p. 153); deltidium is open (see Topper et al. 2013T, fig. 4).
Botsfordia: The homology of the triangular notch or groove in the pseudointerarea with the umbonal pedicle foramen of acrotretids was proposed by Popov (1992), and seemingly corroborated by observations of Ushatinskaya & Korovnikov (2016R), who note that the propareas of the Botsfordia ventral valve sometimes merge to form an elongate teardrop-shaped pedicle foramen.
Clupeafumosus socialis: Following Popov (1992), the larval delthyrium is sealed in adults by outgrowths of the posterolateral margins of the shell.
Eoobolus: See for example fig. 5 in Balthasar 2009T.
Glyptoria: “Delthyrium and notothyrium open, wide” – Cooper 1976.
Longtancunella chengjiangensis: Unclear: a narrow ridge that may correspond to a pseudodeltidium evident in fig 2a and sketched in fig. 2c is not discussed in the text of Zhang et al. 2011T, so the delthyrial region is coded as ambiguous.
Mickwitzia muralensis: A delthyrium is present in young individuals (Balthasar 2004).
Micrina: Opening inferred by Holmer et al. (2008).
Pelagodiscus atlanticus: The listrum (pedicle opening) is interpreted as originating via a similar mechanism to that of acrotretids (Popov 1992), and hence corresponding to a basally sealed delthyrium.
Siphonobolus priscus: Ontogeny presumed to resemble that of acrotretids.
Tonicella: The antemucronal area (Schwabe, 2010) is treated as equivalent to the brachiopod delthyrium.
Yuganotheca elegans: Details of the hinge region are unclear due to the flattened and overprinted nature of fossil preservation.

[157] Posterior surface: Delthyrium: Shape

Character adds 5 to tree score (4 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[2]Micromitra[2]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[2]Terebratulina[2]Orthis[2]Alisina[?]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[2]Gasconsia[3]Pelagodiscus atlanticus[23]Mickwitzia muralensis[1]Mummpikia nuda[2]Botsfordia[2]Eoobolus[3]Lingulellotreta malongensis[-]Lingula[3]Siphonobolus priscus[3]Acanthotretella spinosa[3]Lingulosacculus[3]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 157: Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape

1: Parallel sided2: Triangular3: Round-: InapplicableTransformational character.

A parallel-sided delthyrium links Mummpikia with the Obolellidae (Balthasar 2008).

Following Popov (1992), the larval delthyrium of acrotretids and allied taxa is understood to be sealed in adults by outgrowths of the posterolateral margins of the shell. The resultant round or teardrop-shaped foramen corresponds the delthyrium.

Askepasma toddense: Prominently triangular (see Topper et al. 2013T, fig. 2).
Clupeafumosus socialis: Following the model of Popov (1992).
Mickwitzia muralensis: An opening is incorporated at the base of the homeodeltidium when the organism switches from early to late maturity (fig. 10 in Balthasar 2004). This opening is conceivably homologous with the pedicle foramen of acrotretid brachiopods and their ilk. To reflect this possible homology, Mickwitzia is coded as polymorphic (triangular/round).

[158] Posterior surface: Delthyrium: Shape: Aspect of rounded opening

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[-]Micromitra[-]Askepasma toddense[-]Coolinia pecten[-]Antigonambonites planus[?]Yuganotheca elegans[-]Tomteluva perturbata[-]Salanygolina[-]Terebratulina[-]Orthis[-]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[1]Pelagodiscus atlanticus[3]Mickwitzia muralensis[-]Mummpikia nuda[-]Botsfordia[-]Eoobolus[1]Lingulellotreta malongensis[-]Lingula[2]Siphonobolus priscus[-]Acanthotretella spinosa[2]Lingulosacculus[2]Clupeafumosus socialis[-]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 158: Sclerites: Ventral valve: Posterior surface: Delthyrium: Shape: Aspect of rounded opening

1: Elongate: oval to rhombic2: Essentially circular3: Wider than long-: InapplicableTransformational character.

Chen et al. (2007) propose that an oval to rhombic foramen characterises the discinids (and Heliomedusa, though the foramen in this taxon has since been reinterpreted by Zhang et al. (2009) as an impression of internal tissue).

Lingulellotreta malongensis: Oval (Williams et al. 2000).
Mickwitzia muralensis: Wider than long: see fig. 10 in Balthasar 2004.

[159] Posterior surface: Delthyrium: Cover

Character adds 5 to tree score (4 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[2]Micromitra[1]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[2]Tomteluva perturbata[2]Salanygolina[2]Terebratulina[1]Orthis[2]Alisina[?]Longtancunella chengjiangensis[1]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[2]Gasconsia[-]Pelagodiscus atlanticus[2]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[2]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 159: Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover

1: Open2: Covered, at least in part-: InapplicableTransformational character.

An open delthyrium links Mummpikia with the Obolellidae (Balthasar 2008).

The delthyrial opening can be covered by one or more deltidial plates, or a pseudodeltitium.

Inapplicable in taxa with a round delthiruym (generated by overgrowth of the delthyrial opening by posterolateral parts of the shell, per Popov 1992).

Askepasma toddense: Open (Topper et al. 2013T).
Botsfordia: See pl. 3 fig. 15 in Skovsted & Holmer (2005).
Coolinia pecten: A convex pseudodeltidium completely covers the delthyrium in Coolinia.
Glyptoria: Coded as open by Williams et al. (1998T).
Micromitra: Williams et al. 2000, fig. 83.3.
Nisusia sulcata: “Covered only apically by a small convex pseudodeltitium” – Holmer et al. 2018E.
Paterimitra: Covered by subaical flange, in part.

[160] Posterior surface: Delthyrium: Cover: Extent

Character adds 6 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[2]Micromitra[-]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[2]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[-]Orthis[2]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[1]Nisusia sulcata[2]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[2]Gasconsia[-]Pelagodiscus atlanticus[2]Mickwitzia muralensis[-]Mummpikia nuda[-]Botsfordia[1]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[1]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 160: Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Extent

1: Covered only partially; partially open2: Completely covered?: Ambiguous-: InapplicableTransformational character.

Micrina: Remains somewhat open.
Micromitra: Completely covered (Williams et al. 2000, fig. 83.3).
Nisusia sulcata: A well-defined pseudo-deltidium […] closes only the apical part of
the delthyrium (Rowell & Caruso 1985).

[161] Posterior surface: Delthyrium: Cover: Identity

Character adds 3 to tree score1 additional regionAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[3]Tonicella[3]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[1]Micromitra[-]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[2]Tomteluva perturbata[1]Salanygolina[2]Terebratulina[-]Orthis[12]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[1]Gasconsia[-]Pelagodiscus atlanticus[1]Mickwitzia muralensis[-]Mummpikia nuda[-]Botsfordia[1]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[1]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 161: Sclerites: Ventral valve: Posterior surface: Delthyrium: Cover: Identity

1: Pseudodeltidium2: Deltidial plate(s)3: Continuation of shell?: Ambiguous-: InapplicableTransformational character.

This character has the capacity for further resolution (one or more deltidial plates), but this is unlikely to affect the results of the present study.

The pseudodelthyrium is also referred to as a homeodeltidium.

The antemucronal area of Polyplacophora is treated as equivalent to the brachiopod delthyrium, but is not depositionally distinct to the rest of the shell, so is coded with a distinct character state.

Alisina: Stated as “concave pseudodeltidium with median plication” – Williams et al. 2000
Coded as “Pseudodeltidium: Covered by concave plate” by Bassett et al. (2001).
Askepasma toddense: No pseudodeltidium (Williams et al. 2000, p. 153).
Gasconsia: A homeodeltidium is illustrated by Hanken & Harper (1985).
Lingulellotreta malongensis: The subapical flange of the Paterimitra S1 sclerite has been homologised with the ventral homeodeltidium of Micromitra (Larsson et al 2014).
Mickwitzia muralensis: Termed a homoedeltidium by Balthasar (2004).
Micrina: “Ventral valve convex with apsacline interarea bearing delthyrium, covered by a convex pseudodeltidium” – Holmer et al. 2008.

[162] Posterior surface: Delthyrium: Pseudodeltidium: Shape

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[-]Cupitheca holocyclata[-]Bactrotheca[-]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[2]Micromitra[-]Askepasma toddense[2]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[2]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[-]Orthis[1]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[2]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[1]Gasconsia[-]Pelagodiscus atlanticus[2]Mickwitzia muralensis[-]Mummpikia nuda[-]Botsfordia[1]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 162: Sclerites: Ventral valve: Posterior surface: Delthyrium: Pseudodeltidium: Shape

1: Concave2: Convex?: Ambiguous-: InapplicableTransformational character.

A ridge-like (i.e. convex) pseudodeltitium unites Salanygolina with Coolinia and other Chileata (Holmer et al. 2009, p. 6).

Alisina: “concave pseudodeltidium with median plication” – Williams et al. 2000
Coded as “Pseudodeltidium: Covered by concave plate” by Bassett et al. (2001).
Antigonambonites planus: Convex (Williams et al. 2000, fig. 508).
Gasconsia: “Narrow depressed homeodeltidium” – Hanken & Harper (1985).
Kutorgina chengjiangensis: Difficult to determine based on material presented in Zhang et al (2007R), or indeed for other species in the genus (e.g. Williams et al. 2000; Skovsted & Holmer 2005; Holmer et al. 2018T).
Mickwitzia muralensis: Convex (see Balthasar 2004, fig. 4B).
Micrina: Convex deltoid (Holmer et al. 2008T).
Micromitra: Gently convex (see Williams et al. 2000, fig. 83.3).
Nisusia sulcata: Convex in Nisusia (see Rowell and Caruso, 1985, fig. 8.4).
Paterimitra: Gently convex (see Williams et al. 2000, fig. 83.1).
Salanygolina: “The presence of […] a narrow delthyrium covered by a convex pseudodeltidium, places Salanygolinidae outside the Class Paterinata and strongly suggests affinity to the Cambrian Chileida” – Holmer et al. 2009, p. 9.
Tomteluva perturbata: Convex (Streng et al. 2016).

[163] Posterior surface: Delthyrium: Pseudodeltidium: Hinge furrows

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[?]Longtancunella chengjiangensis[0]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 163: Sclerites: Ventral valve: Posterior surface: Delthyrium: Pseudodeltidium: Hinge furrows

0: Absent1: PresentNeomorphic character.

After Bassett et al. (2001) character 18, “Hinge furrows on lateral sides of pseudodeltidium”.

Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina, Phoronis, Dailyatia, Acanthotretella spinosa, Askepasma toddense, Micromitra, Clupeafumosus socialis, Eccentrotheca, Heliomedusa orienta, Lingulosacculus, Lingulellotreta malongensis, Micrina, Mummpikia nuda, Orthis, Paterimitra, Bactrotheca, Haplophrentis carinatus, Pedunculotheca diania: Absent due to inapplicability of neomorphic character.
Gasconsia: Not evident or illustrated (Hanken & Harper, 1985).
Glyptoria: Coded as absent in Bassett et al. 2001 (table 18.1).
Kutorgina chengjiangensis, Nisusia sulcata: Coded as present in Bassett et al. 2001 (table 18.1).
Salanygolina: The presence of this feature is impossible to determine based on the available material.

[164] Umbonal perforation

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[1]Cupitheca holocyclata[?]Bactrotheca[1]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[0]Terebratulina[0]Orthis[1]Alisina[1]Longtancunella chengjiangensis[0]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[?]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 164: Sclerites: Ventral valve: Umbonal perforation

0: Umbo imperforate (or ventral valve absent)1: Umbonal perforation?: AmbiguousNeomorphic character.

Certain taxa, particularly those with a colleplax, exhibit a perforation at the umbo of the ventral valve. This opening is sometimes associated with a pedicle sheath, which emerges from the umbo of the ventral valve without any indication of a relationship with the hinge.

In contrast, the pedicle of acrotretids and similar brachiopods is situated on the larval hinge line, but is later surrounded by the posterolateral regions of the growing shell to become separated from the hinge line, and encapsulated in a position close to (or with resorption of the brephic shell, at) the umbo (see Popov 1992, pp. 407–411 and fig. 3 for discussion). In some cases, an internal pedicle tube attests to this origin – potentially corresponding to the pedicle groove of lingulids. As such, the pedicle foramen of acrotretids and allies is not originally situated at the umbo; it is instead understood to represent a basally sealed delthyrium.

Bactrotheca: The apical termination of the conical valve is not preserved (Valent et al., 2012).
Clupeafumosus socialis: The presumed pedicle foramen reported by Topper et al. (2013R) is at the ventral valve umbo. No evidence of an internal pedicle tube is present, but we follow Popov (1992) in inferring the encapsulation of the pedicle foramen.
Cupitheca holocyclata: Decollation generates open umbo, sealed secondarily with septum.

Dailyatia: The B and C sclerites of Dailyatia bear small umbonal perforations (Skovsted et al 2015), but these are not considered to be homologous with the ventral valve, so this character is coded as inapplicable – though the possibility that the perforations are equivalent is intriguing.

A1 sclerites typically have a pair of perforations, which are conceivably equivalent to the setal tubes of Micrina (Holmer et al. 2011). The A1 sclerite of D. bacata has a structure that is arguably similar to the ‘colleplax’ of Paterimitra. But the homology of any of these structures to the umbonal aperture of brachiopods is difficult to establish.
Eccentrotheca: The sclerites of Eccentrotheca form a ring that surrounds the inferred attachment structure; the attachment structure does not emerge from an aperture within an individual sclerite. Thus no feature in Eccentrotheca is judged to be potentially homologous with the apical perforation in bivalved brachiopods.
Heliomedusa orienta: There is “compelling evidence to demonstrate that the putative pedicle
illustrated by Chen et al. (2007: Figs. 4, 6, 7) in fact is the mold of a three-dimensionally preserved visceral cavity.” – Zhang et al. 2009.
Lingulosacculus: The apical termination of the fossil is unknown.
Mickwitzia muralensis: The umbo itself is imperforate (Balthasar 2004).
Paramicrocornus: Z.-L. Zhang et al. (2018).
Paterimitra: The presumed pedicle foramen is an opening between the S1 and S2 sclerites, neither of which are perforated (Skovsted et al. 2009).
Siphonobolus priscus: Prominent subcircular perforation at umbo associated with an internal pedicle tube (Popov et al. 2009), thus presumed to share an origin with the acrotretid pedicle foramen.
Tomteluva perturbata: Streng et al (2016) observe “an internal tubular structure probably representing the ventral end of the canal within the posterior wall of the pedicle tube”, but do not consider this tomteluvid dube to be homologous with the pedicle tube of acrotretids and their ilk, stating (p. 274) that it appears to be unique within Brachiopoda.

[165] Umbonal perforation: Shape

Character adds 3 to tree score (2 with Fitch)2 additional regions (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[-]Leptochiton[-]Glaphurochiton carbonarius[-]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[-]Haplophrentis carinatus[3]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[-]Cotyledion tylodes[-]Eccentrotheca[-]Orthrozanclus[-]Micromitra[-]Askepasma toddense[1]Coolinia pecten[2]Antigonambonites planus[?]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[-]Terebratulina[-]Orthis[1]Alisina[1]Longtancunella chengjiangensis[-]Glyptoria[1]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[-]Craniops[-]Ussunia[-]Heliomedusa orienta[-]Gasconsia[-]Pelagodiscus atlanticus[-]Mickwitzia muralensis[-]Mummpikia nuda[-]Botsfordia[-]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[?]Lingulosacculus[12]Clupeafumosus socialis[-]Micrina[-]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 165: Sclerites: Ventral valve: Umbonal perforation: Shape

1: Circular (or subcircular)2: Rhombic to oval3: Arising through decollation?: Ambiguous-: InapplicableTransformational character.

The perforation in Cupitheca seems to have a distinct origin, arising through decollation; as such, the shape simply reflects the outline of the shell. This reflects a distinct origin of the perforation and is therefore provided as a separate state.

Acanthotretella spinosa: Too small to observe given quality of preservation (Holmer & Caron 2006).
Alisina: Seemingly circular (Zhang et al. 2011A).
Antigonambonites planus: Based on p.92, fig.4B.
Clupeafumosus socialis: Taller than wide in some cases, but very nearly circular in others; see Topper et al. (2013R).
Coolinia pecten: Bassett and Popov write “a dominant feature of the ventral umbo is a sub-oval perforation about 270 μm long and 250 μm wide”: the width and height of this structure are almost identical, and we score it as (sub) circular.
Heliomedusa orienta: Rhombic to oval – seen as evidence for a discinid affinity (Chen et al. 2007).
Kutorgina chengjiangensis: The exact size and shape of the apical perforation is obscured by the emerging pedicle.
Nisusia sulcata: “close to circular” (Holmer et al. 2018E).
Salanygolina: Essentially circular (Holmer et al. 2009, fig. 4).

[166] Colleplax, cicatrix or pedicle sheath

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[?]Tomteluva perturbata[1]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[1]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[1]Craniops[0]Ussunia[?]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[1]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 166: Sclerites: Ventral valve: Colleplax, cicatrix or pedicle sheath

0: Absent1: Present?: AmbiguousNeomorphic character.

In certain taxa, the umbo of the ventral valve bears a colleplax, cicatrix or pedicle sheath; Bassett et al. (2008) consider these structures as homologous.

Bactrotheca: The apical termination of the conical valve is not preserved (Valent et al., 2012).
Botsfordia: Following Williams et al. 1998T, appendix 2.
Clupeafumosus socialis: Not reported by Topper et al. (2013R).
Craniops: Paracraniops is “externally similar to Craniops, but lacking cicatrix” – indicating that Craniops bears a cicatrix (Williams et al. 2000). Also coded as present in their table 15.
Heliomedusa orienta: A cicatrix was reconstructed by Jin & Wang 1992 (figs 6b, 7), but has not been reported by later authors; possibly, as with the ‘pedicle foramen’ of Chen et al. (2007), this structure represents internal organs rather than a cicatrix proper (Zhang et al. 2009); as such it has been recorded as ambiguous.
Kutorgina chengjiangensis: The umbonal region of kutorginides “clearly lacks a pedicle sheath” (Holmer et al. 2018T).
Lingulellotreta malongensis: The pedicle is identified as such (rather than a pedicle sheath) by the internal pedicle tube.
Longtancunella chengjiangensis: A ring-like structure surrounding the pedicle is interpreted as a colleplax (Zhang et al. 2011T), though the authors make no comparison with the pedicle capsule exhibited by extant terebratulids (see Holmer et al. 2018E).
Micrina: Absent in Micrina (Holmer et al. 2011).
Pedunculotheca diania: The flat apical termination of juvenile individuals possibly functioned as colleplax for attachment, but may simply represent the brephic shell; we treat it as ambiguous to reflect this potential homology.
Siphonobolus priscus: Coded as present in view of the attachment scar, which has been considered homologous with the “adult colleplax and foramen with attachment pad” in Salanygolina (Popov et al. 2009).
Tomteluva perturbata: The internal canal associated with the pedicle is unique to the tomteluvids, and has an uncertain identity (Streng et al. 2016). It could conceivably correspond to an internalized pedicle sheath or an equivalent structure, so this feature is coded as ambiguous here.
Ussunia: Following table 15 in Williams et al. 2000.
Yuganotheca elegans: The median collar or conical tube is conceivably homologous with the pedicle sheath.

[167] Median septum

Character adds 6 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[0]Askepasma toddense[0]Coolinia pecten[1]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[0]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[0]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 167: Sclerites: Ventral valve: Median septum

0: Absent1: PresentNeomorphic character.

Chen et al. (2007) observe a median septum in what they interpret as the ventral valve of Heliomedusa, and the ventral valve of Discinisca, which they propose points to a close relationship.

Acanthotretella spinosa: Carbonaceous preservation confounds the identification of internal shell structures; it is possible that this feature is present, but not observable in the Burgess Shale material.
Botsfordia: Following Williams et al. 1998T, appendix 2.
Clupeafumosus socialis: A short medial ridge (septum) is present in the ventral valve (Topper et al. 2013R).
Eoobolus: Prominent median septum (fig. 4d, e in Balthasar 2009T).
Gasconsia: Evident in moulds of ventral valve (Hanken & Harper, 1985; Watkins, 2002).
Glyptoria: Neither evident nor reported in Williams et al. (2000).
Haplophrentis carinatus: The carina of H. carinatus is an angular elevation of the ventral valve surface, rather than a septum growing inward on the interior of shell.
Heliomedusa orienta: Reported on ‘ventral’ valve by Chen et al. (2007); we consider the ‘ventral’ valve to be the dorsal valve.
Lingulellotreta malongensis: Medial septum visible in ventral valve in Williams et al. (2000), fig. 34.1c.
Micromitra: Ventral ridge characteristic of Micromitra (Skovsted & Peel 2010).
Mummpikia nuda: “Some specimens also reveal that the vault had a slight median septum, which is now visible as a notch or a groove dividing the right from the left part” – Balthasar 2008.
Novocrania: Valve thin and often unmineralized.
Pelagodiscus atlanticus: Described as present in Discinisca by Chen et al. 2007; assumed present also in Pelagodiscus.
Siphogonuchites multa: Seemingly present in S. multa (Bengtson, 1992, fig. 2), though this is not interpreted as a ventral valve.
Siphonobolus priscus: Present; see Popov et al. 2009, fig. 5J.
Ussunia: Following char. 42 in table 15 in Williams et al. 2000.

4.16 Sclerites: Ornament

[168] Concentric ornament

Character adds 13 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[2]Leptochiton[2]Glaphurochiton carbonarius[2]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[2]Neopilina[2]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[1]Dentalium[1]Pelagiella[1]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[1]Paramicrocornus[2]Haplophrentis carinatus[2]Cupitheca holocyclata[1]Bactrotheca[2]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[2]Dailyatia[2]Cotyledion tylodes[2]Eccentrotheca[2]Orthrozanclus[2]Micromitra[2]Askepasma toddense[1]Coolinia pecten[2]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[2]Salanygolina[1]Terebratulina[1]Orthis[2]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[2]Novocrania[2]Craniops[1]Ussunia[2]Heliomedusa orienta[2]Gasconsia[1]Pelagodiscus atlanticus[2]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[12]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[2]Micrina[2]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 168: Sclerites: Ornament: Concentric ornament

1: Smooth, or growth lines only2: Concentric ornament present?: Ambiguous-: InapplicableTransformational character.

After character 11 in Williams et al. (1998T). Coded as transformational as it is possible that maintaining a smooth shell without occasional prominent ridges requires greater secretory control.

Acaenoplax hayae: None evident (???).
Askepasma toddense, Micromitra, Glyptoria, Kutorgina chengjiangensis, Salanygolina: Following appendix 2 in Williams et al. (1998T).
Bactrotheca: Both valves with fine growth lines only (Valent et al., 2012).
Botsfordia: Following Williams et al. 1998T, appendix 2.
Pustules are arranged along concentric growth lines (Skovsted & Holmer, 2005), so are not treated as a distinct ornamentation.
Calvapilosa kroegeri: Prominent ridge in certain specimens (Vinther et al., 2017).
Conocardium elongatum: Some concentric ornament evident in some regions of the shell (Rogalla & Amler, 2003).
Cotyledion tylodes: Zhang et al. (2013).
Cupitheca holocyclata: Broad symmetric ridges in some specimens (Vendrasco et al., 2017).
Eccentrotheca: More or less concentric ridges occur on Eccentrotheca sclerites (Skovsted et al. 2011).
Glaphurochiton carbonarius: Concentric ridges, with additional pustules (Hoare & Mapes, 1986).
Halkieria evangelista: Ridges in shell parallel, but are more prominent than, growth lines.
Haplophrentis carinatus: A series of regularly spaced concentric ridges adorn both valves (Moysiuk et al. 2017); these are more pronounced than mere growth lines.
Heliomedusa orienta: The ornament on shell exterior is described as concentric fila (Chen et al., 2007, P.43), and also scored as it in Williams et al. (2000, pp.160–163).
Leptochiton: (???).
Mickwitzia muralensis: Symmetric fila.
Mytilus: Growth lines only.
Neopilina: Menzies & Layton (1962).
Novocrania: Irregular ridges externally (Williams et al. 2000).
Orthrozanclus: Concentric ridges in addition to growth lines (Conway Morris & Caron, 2007).
Pedunculotheca diania: A series of regularly spaced concentric ridges adorn the ventral valve; comparatively less regular lines ornament the operculum.
Pelagiella: Ornament, if present, is not concentric.
Pelagodiscus atlanticus: Only growth lines evident (Williams et al. 2000).
Phthipodochiton thraivensis: “Ornament of growth lines only” (Sutton & Sigwart, 2012).
Terebratulina: Single ridge evident in Williams et al. (2006) fig. 1425.1a interpreted as interruption ot growth rather than inherent feature, so coded as absent (i.e. smooth).
Tonicella: No prominent ornamentat in Tonicella (Connors et al., 2012).
Yilingia spiciformis: Smooth (Chen et al., 2019).

[169] Concentric ornament: Symmetry

Character adds 8 to tree score (6 with Fitch)3 additional regions (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[-]Mopalia[-]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[2]Neopilina[2]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[-]Paramicrocornus[2]Haplophrentis carinatus[2]Cupitheca holocyclata[-]Bactrotheca[2]Pedunculotheca diania[1]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[1]Dailyatia[1]Cotyledion tylodes[1]Eccentrotheca[?]Orthrozanclus[1]Micromitra[2]Askepasma toddense[-]Coolinia pecten[1]Antigonambonites planus[-]Yuganotheca elegans[-]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[-]Orthis[1]Alisina[-]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[-]Ussunia[2]Heliomedusa orienta[1]Gasconsia[-]Pelagodiscus atlanticus[2]Mickwitzia muralensis[-]Mummpikia nuda[-]Botsfordia[-]Eoobolus[-]Lingulellotreta malongensis[-]Lingula[-]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[-]Clupeafumosus socialis[12]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 169: Sclerites: Ornament: Concentric ornament: Symmetry

1: Asymmetric fila, with outer faces2: Symmetric fila?: Ambiguous-: InapplicableTransformational character.

After character 11 in Williams et al. (1998T).

Alisina: Seemingly asymmetric (Williams et al. 2000, fig. 122.3c; Zhang et al. 2011A, Fig. 1).
Askepasma toddense, Micromitra, Glyptoria, Kutorgina chengjiangensis, Salanygolina: Following appendix 2 in Williams et al. (1998T).
Calvapilosa kroegeri: Vinther et al. (2017).
Dailyatia: Clear asymmetry (Skovsted et al. 2015).
Eccentrotheca: Ornament, such as it is, is asymmetric, with prominent outer faces (Skovsted et al. 2011).
Gasconsia: Assymmetric (Hanken & Harper, 1985, fig. 3).
Heliomedusa orienta: See fig. 1715 in Williams et al. (2007).
Leptochiton: Kaas (1981).
Mickwitzia muralensis: Symmetric fila (Balthasar 2004).
Micrina: No obvious asymmetry, even if not obviously symmetric either (Holmer et al. 2008). Coded as ambiguous.
Neopilina: Seemingly symmetric (Menzies & Layton, 1962).
Novocrania: Clear outer faces (Williams et al. 2000, fig. 100.2b).
Orthrozanclus: Preservation inadequate to determine.

[170] Radial ornament

Character adds 12 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[1]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[1]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[1]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[0]Yuganotheca elegans[1]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[1]Orthis[0]Alisina[0]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[1]Gasconsia[0]Pelagodiscus atlanticus[1]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[1]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 170: Sclerites: Ornament: Radial ornament

0: Absent1: Present?: AmbiguousNeomorphic character.

Ridges radiating from umbo, i.e. ribs.

Askepasma toddense: “Ornament of irregularly developed, concentric growth lamellae; microornament of irregularly arranged, polygonal pits” – Williams et al. 2000, p153; figs on p.155.
Botsfordia: Following Williams et al. 1998T, Appendix 2.
Calvapilosa kroegeri: Radial structures interpreted as aesthete canals (Vinther et al., 2017).
Eoobolus: Very faint costellae in some specimens but coded absent.
Gasconsia: “Ornament of indistinct low radial ribs” – Williams et al. (2000, p167).
Glyptoria: “Coarsely costate” – Williams et al. (2000, p710).
Haliotis: Radial (apical to apertural) lineations present (Auzoux-Bordenave et al., 2010).
Heliomedusa orienta: See fig. 1715 in Williams et al. (2007).
Neopilina: Menzies & Layton (1962).
Orthrozanclus: Conway Morris & Caron (2007).
Paramicrocornus: Z.-L. Zhang et al. (2018).
Pelagiella: Radial arrangement of knobs (???).
Polysacos vickersianum: Radial ridges present on certain valves; particularly evident on tail valve (???).
Siphonobolus priscus: “Indistinct radial ribs accentuated by radial rows of tubercles” – Popov et al. 2009.
Ussunia: Unornamented.
Yilingia spiciformis: Smooth (Chen et al., 2019).

[171] Shell-penetrating spines

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[1]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 171: Sclerites: Ornament: Shell-penetrating spines

0: Absent1: PresentNeomorphic character.

Mineralized or partly mineralized spines are observed in Heliomedusa and Acanthotretella.

Acaenoplax hayae: Spines are processes of the shells, rather than penetrative (???).
Glyptoria: Neither evident nor reported in Williams et al. (2000).
Heliomedusa orienta: The ‘spines’ reported by Chen et al. (2007) are pyritized spinelike setae – see pp. 2580–2590 in Williams et al. (2007).
Nisusia sulcata: Bears numerous small, hollow spines (Williams et al. 2000).
Orthrozanclus: Not evident (Conway Morris & Caron, 2007).
Siphogonuchites multa: No treated as homologous to those of brachiopods, due to their inferred homology with setae.
Tonicella: Aesthete canals penetrate the main valves of certain chitons, but are not equivalent to the shell-penetrating spines of brachiopods.

4.17 Sclerites: Composition

[172] Mineralogy

Character adds 12 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[4]Tonicella[4]Mopalia[3]Polysacos vickersianum[4]Leptochiton[34]Glaphurochiton carbonarius[34]Calvapilosa kroegeri[34]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[4]Neopilina[4]Conocardium elongatum[4]Haliotis[4]Pojetaia runnegari[4]Mytilus[-]Dentalium[4]Pelagiella[4]Siphogonuchites multa[4]Acaenoplax hayae[34]Kulindroplax perissokomos[4]Paramicrocornus[4]Haplophrentis carinatus[4]Cupitheca holocyclata[4]Bactrotheca[4]Pedunculotheca diania[4]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[3]Namacalathus[-]Phoronis[2]Dailyatia[234]Cotyledion tylodes[2]Eccentrotheca[34]Orthrozanclus[2]Micromitra[2]Askepasma toddense[3]Coolinia pecten[3]Antigonambonites planus[1]Yuganotheca elegans[3]Tomteluva perturbata[23]Salanygolina[3]Terebratulina[3]Orthis[3]Alisina[?]Longtancunella chengjiangensis[3]Glyptoria[3]Nisusia sulcata[3]Kutorgina chengjiangensis[3]Novocrania[3]Craniops[4]Ussunia[12]Heliomedusa orienta[4]Gasconsia[2]Pelagodiscus atlanticus[2]Mickwitzia muralensis[3]Mummpikia nuda[2]Botsfordia[23]Eoobolus[2]Lingulellotreta malongensis[2]Lingula[2]Siphonobolus priscus[1234]Acanthotretella spinosa[14]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[2]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 172: Sclerites: Composition: Mineralogy

1: Organic (non-mineralized)2: Phosphatic3: Calcitic4: Aragonitic?: Ambiguous-: InapplicableTransformational character.

Acaenoplax hayae: Preserved as calcite, but interpreted as aragonitic (???).

Acanthotretella spinosa: Holmer & Caron (2006) note the absence of brittle breakage, interpreted as indicating the absence of a material mineralized component to the shells. The preservation is strikingly different from that of other Burgess Shale brachiopods, ruling out a primarily calcitic or phosphatic composition. The two-dimensional nature of the preservation also differs from that of co-occurring aragonitic taxa (hyoliths; Holmer & Caron 2006 p. 273), indicating that any mineralization was minor at best.

Holmer & Caron (2006, p. 286) suggest that it is more likely that a (minor) mineral component was present than that it was not, though without providing an uncontestable rationale. To be as conservative as possible, we therefore code this taxon as ambiguous.
Calvapilosa kroegeri: Calcareous (Vinther et al., 2017).
Clupeafumosus socialis: Phosphatic – hence the conventional placement within Linguliformea.
Conocardium elongatum: Entirely aragonitic in Apotocardium (Rogalla, Carter, & Pojeta, 2003).
Cotyledion tylodes: The extensive relief and association with pyrite framboids indicates original mineralization, but the identity of the biomineral remains uncertain (Zhang et al., 2013).
Craniops: Shell calcitic.
Cupitheca holocyclata: Reconstructed as aragonitic from microstructural fabrics (Vendrasco et al., 2017).
Eoobolus: “the original shell of Eoobolus contained small calcareous grains that were incorporated into organic-rich layers alongside apatite” (Balthasar 2007).
Gasconsia: Confirmed in Trimerella by Balthasar et al. 2011.
Haliotis: “Essentially made of aragonite” (Auzoux-Bordenave et al., 2010).
Heliomedusa orienta: “Shell originally organophosphatic, but may generally have been poorly mineralized” – Williams et al. 2007 – cf. ibid, p. 2889, “These strong similarities to discinoids in soft-part anatomy imply that the Heliomedusa shell was chitinous or chitinophosphatic, not calcareous.”
Kulindroplax perissokomos: Presumed calcareous.
Lingulellotreta malongensis: Coded as phosphatic by Zhang et al. (2014), but with no explanation.
Cracks within shells of Chengjiang specimens (e.g. Zhang et al. 2007N, fig. 3) demonstrate that the shells were originally mineralized, but not the identity of the original biomineral. This said, phosphatized material from Kazakhstan (Holmer et al. 1997) is attributed to the same species; presuming this phosphate to be original and the material to be conspecific, L. malongensis is coded as having phosphatic shells.
Lingulosacculus: The absence of relief in Lingulosacculus rules out a phosphatic or calcitic composition, but co-occurring (and presumably aragonitic) hyolithids are preserved in the same fashion. Its constitution was thus either organic or aragonitic (Balthasar & Butterfield 2009E).
Longtancunella chengjiangensis: “The original composition of the shell cannot be determined with certainty”, though it was “most probably entirely soft and organic” – Zhang et al. 2011T.
Mickwitzia muralensis: Calcite and silica deemed diagenetic by Balthasar (2004).
Mummpikia nuda: Identified as calcareous by preservational criteria, and description “primary
calcitic shells of M. nuda” (Balthasar 2008).
Novocrania: Ventral valve uncalcified in extant forms or sometimes thin (Williams et al., 2000), but coded as calcitic as calcite-mineralizing pathways are present.
Orthrozanclus: Relief indicates original mineralization, presumably in calcium carbonate as the originally phosphatic biominerals retain their original composition in Burgess Shale palaeoscolecids (???).
Pelagiella: Aragonite (???).
Pojetaia runnegari: Originally comprised spherulitic aragonite prisms (???).
Polysacos vickersianum: By analogy with close relative Protobalanus (Vinther et al., 2012).
Salanygolina: Original mineralogy unknown, but known to be mineralised and anticipated to be phosphatic (Holmer et al. 2009).
Siphogonuchites multa: Interpreted as aragonitic (Bengtson, 1992).
Ussunia: Trimerellids were probably aragonitic (Williams et al., 2000).
Yilingia spiciformis: Presumably non-mineralized, on account on the absence of equivalent features in the shelly fossil record.

[173] Cuticle or organic matrix

Character adds 3 to tree score (2 with Fitch)1 additional region (not counted by Fitch)An optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[-]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 173: Sclerites: Composition: Cuticle or organic matrix

1: GAGs, chitin and collagen2: Glycoprotein?: Ambiguous-: InapplicableTransformational character.

Williams et al. (1996) identify glycoprotein-based organic scaffolds as distinct from those comprising glycosaminoglycans (GAGs), chitin and collagen. This character can only be scored for extant taxa.

Haliotis: “The organic matrix [..] is a mixture of proteins, glycoproteins, lipids, chitin and acidic polysaccharides” (Auzoux-Bordenave et al., 2010) – no GAGs or collagen.
Lingula: Coded as GAGs, chitin and collagen in lingulids by Williams et al (1996).
Mytilus: Glycolytic domain containing proteins were observed (=glycoproteins?), whereas GAGs were not (Gao et al., 2015).
Novocrania: Coded as glycoprotein for craniids by Williams et al (1996).
Pelagodiscus atlanticus: Coded as GAGs, chitin and collagen in discinids by Williams et al (1996).
Phoronis: “The presence of sulphated glycosaminoglycans (GAGs) in the chitinous cuticle of Phoronis (Herrmann, 1997, p. 215) would suggest a link with linguliforms, as GAGs are unknown in rhynchonelliform shells (Fig. 1891, 1896)” – Williams et al. 2007, p. 2830.
Siphonobolus priscus: Lenticular chambers in siphonotretid shells interpreted as degraded GAG residue (Williams et al. 2004).
Terebratulina: Coded as glycoprotein for terebratulids by Williams et al (1996).

[174] Incorporation of sedimentary particles

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[1]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[1]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 174: Sclerites: Composition: Incorporation of sedimentary particles

0: Absent1: PresentNeomorphic character.

Phoronids and Yuganotheca aggulutinate particles into their sclerites.

[175] Microstructure: Number of distinct layers

Character adds 9 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[4]Tonicella[?]Mopalia[?]Polysacos vickersianum[4]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[2]Neopilina[2]Conocardium elongatum[4]Haliotis[1]Pojetaia runnegari[4]Mytilus[-]Dentalium[4]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[2]Paramicrocornus[?]Haplophrentis carinatus[2]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[1]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[3]Namacalathus[-]Phoronis[1]Dailyatia[?]Cotyledion tylodes[1]Eccentrotheca[?]Orthrozanclus[3]Micromitra[3]Askepasma toddense[2]Coolinia pecten[4]Antigonambonites planus[-]Yuganotheca elegans[?]Tomteluva perturbata[2]Salanygolina[2]Terebratulina[2]Orthis[?]Alisina[?]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[2]Novocrania[2]Craniops[?]Ussunia[?]Heliomedusa orienta[2]Gasconsia[2]Pelagodiscus atlanticus[2]Mickwitzia muralensis[1]Mummpikia nuda[2]Botsfordia[2]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[2]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 175: Sclerites: Composition: Microstructure: Number of distinct layers

1: Single microstructural layer2: Two microstructurally differentiated layers3: Inner and outer laminae enclosing medial void4: Three microstrurally differentiated layers?: Ambiguous-: InapplicableTransformational character.

Hyolith conchs comprise two mineralized layers of fibrous bundles. Bundles are measure 5–15 μm across; their constituent fibres are each 0.1–1.0 μm wide. In the inner layer, the fibres are transverse; in the outer layer, the bundles are inclined towards the umbo, becoming longitudinal on the outermost margin.

Coded as non-additive as there is no clear necessity to add layers sequentially: for example, three layers could arise by the addition of a void within a single pre-existing layer.

Stratiform laminae, shell-penetrating canals and other features above the scale of crystal organization are not considered as contributing to the mineralogical microstructure and are coded separately.

Inapplicable in taxa with a non-mineralized shell.

Bactrotheca: Taxon known only from moulds (Valent et al., 2012).
Botsfordia: “Composed of a thin primary layer and a laminate secondary shell exhibiting baculate shell structure” – Skovsted & Holmer (2005), with reference to Skovsted & Holmer 2003.
Clupeafumosus socialis: General acrotretid structure taken from Zhang et al. (2016).
Conocardium elongatum: Two layers are microstructrually differentiated; the ‘inner’ layer is considered a sub-layer of the ‘middle’ layer (Rogalla et al., 2003).
Cupitheca holocyclata: Inner and outer layers (Vendrasco et al., 2017).
Eoobolus: “Eoobolus shells exhibit the general characteristics of modern linguliform shells, i.e. they were composed of alternating sets of organic and apatite-rich layers that were separated by thin sheets of recalcitrant organic layers.” – Balthasar 2007.
Haliotis: (In juveniles): tablets plus inner and outer prismatic layers (Auzoux-Bordenave et al., 2010).
Halkieria evangelista: Single layer of fibrous aragonite (Porter, 2008).
Leptochiton: (???).
Mickwitzia muralensis: “the shell structure of Mickwitzia […] is closely similar to the columnar shell of linguliform acrotretoid brachiopods as well as to the linguloid Lingulellotreta, in that it has slender columns in the laminar succession” – Williams et al. 2007.
Micrina: Identical to Mickwitzia and more derived linguliforms (Holmer et al 2011).
Mummpikia nuda: Balthasar (2008) considers the single mineralogical layer, which comprises phosphatic rods and laminae, to characterize Obolellata.
Mytilus: Aragonitic nacre, fibrous calcite prisms, and myostracum (Gao et al., 2015).
Namacalathus: Namacalathus exhibits three layers, none of which have any obvious correspondence with those of brachiopods.
Neopilina: “Shell layers consisting of a thin periostracum, a dominant prismatic layer, and a thin internal nacreous layer” (McLean, 1979)
“Spherulitic aragonitic prisms beneath the organic periostracum” (???).
Paramicrocornus: “The shells of both skeletal components show very similar structures and are composed of two layers” – Z.-L. Zhang et al. (2018).
Pelagiella: At least three microstructures are evident, although it is not quite apparent whether these occur in separate layers or separate regions of the shell (???).
Siphonobolus priscus: “Orthodoxly secreted primary and secondary layers” – Williams et al. 2004.
Tonicella: From periostracum inwards, Chiton bears three microstructural layers: fine-grained, nacreous, and regular crossed lamellar.

[176] Microstructure: Format

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[3]Tonicella[?]Mopalia[?]Polysacos vickersianum[34]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[34]Neopilina[34]Conocardium elongatum[4]Haliotis[4]Pojetaia runnegari[34]Mytilus[-]Dentalium[12]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[2]Paramicrocornus[?]Haplophrentis carinatus[2]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[2]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[1]Namacalathus[-]Phoronis[1]Dailyatia[?]Cotyledion tylodes[1]Eccentrotheca[?]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[-]Yuganotheca elegans[?]Tomteluva perturbata[1]Salanygolina[2]Terebratulina[2]Orthis[?]Alisina[?]Longtancunella chengjiangensis[2]Glyptoria[2]Nisusia sulcata[2]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[?]Ussunia[?]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 176: Sclerites: Composition: Microstructure: Format

1: Laminated2: Fibrous bundles3: Nacreous / crossed lamellar4: Spherulitic prisms?: Ambiguous-: InapplicableTransformational character.

Hyolith conchs comprise two mineralized layers of fibrous bundles. Bundles measure 5–15 μm across; their constituent fibres are each 0.1–1.0 μm wide. In the inner layer, the fibres are transverse; in the outer layer, the bundles are inclined towards the umbo, becoming longitudinal on the outermost margin.

Stratiform laminae, shell-penetrating canals and other features above the scale of crystal organization are not considered as contributing to the mineralogical microstructure and are coded separately.

The pervasive (not just superficial) polygonal structures in Paterimitra are distinct, and characterize Askepasma, Salanygolina, Eccentrotheca and Paterimitra (Larsson et al. 2014)

Williams et al. (2000) identify cross-bladed laminae as diagnostic of Strophomenata, with the exception of some older groups that contain fibres or laminar laths.

Antigonambonites planus: Tabular laminae, not fibrous as previously thought (Madison, 2017).
Askepasma toddense: Lamination present (Balthasar, Skovsted, Holmer, & Brock, 2009), with imprints of presumed mantle cells (following Williams et al., 1998b, appendix 2).

Botsfordia: “Composed of a thin primary layer and a laminate secondary shell exhibiting baculate shell structure” – Skovsted & Holmer (2005), with reference to Skovsted & Holmer 2003.

Williams et al. (1998b appendix 2) code lamination as present, with no imprints of presumed mantle cells.
Conocardium elongatum: Spherulitic prisms are present in the outer layer of Apotocardium; crossed lamellae or prisms in the inner (Rogalla et al., 2003).
Coolinia pecten: Dewing (2004).
Craniops: (Williams et al., 1997, fig. 249.1).
Cupitheca holocyclata: Fibrous bundles (Vendrasco et al., 2017).
Eccentrotheca, Paterimitra: Laminated (Balthasar et al., 2009).
Micromitra, Eoobolus: Lamination present, with no imprints of presumed mantle cells (following Williams et al., 1998b, appendix 2).
Gasconsia: Laminated relict shell structure visible, indicating original constitution from “sheet-like laminae” (Hanken & Harper, 1985).
Glyptoria, Kutorgina chengjiangensis: Lamination absent (following Williams et al., 1998b, appendix 2).
Haliotis: (prismatic) (Auzoux-Bordenave et al., 2010).
Leptochiton: Crossed lamellar sandwiching spherulitic (???).
Lingula: Lingulid laminae are thicker than those of tommotiids or paterinids, but construed as homologous (Balthasar et al., 2009).
Mickwitzia muralensis: Alternation of layers (Balthasar, 2004).
Micrina: Micrina exhibits polygonal imprints on the internal surfaces of successive second-order laminae, suggesting the existence of a polygonal organization of these layers (Balthasar et al., 2009).
Mummpikia nuda: Balthasar (2008) considers the single mineralogical layer, which comprises phosphatic rods and laminae, to characterize Obolellata.
Namacalathus: The inner and outer layer are foliated. The columnar inflections lack canals, and as such we do not consider them to bear any obvious homology with the hollow pillars of tommotiids and certain brachiopods, their superficial similarity to strophomenid pseudopunctae notwithstanding.
Neopilina: “Shell layers consisting of a thin periostracum, a dominant prismatic layer, and a thin internal nacreous layer” (McLean, 1979).
Novocrania: Laminar secondary layer (Parkinson, Curry, Cusack, & Fallick, 2005).
Orthis: Orithidina have impunctate shells with a fibrous secondary layer (Williams et al. 2000, p. 724).
Paramicrocornus: Fibrous (Z.-L. Zhang et al., 2018).
Pojetaia runnegari: Spherulitic aragonitic prisms (???).
Salanygolina: “Apatitic crystals of varying shapes and dimensions” – Holmer et al. (2009).
Siphonobolus priscus: Cleaved stratified platy laminae, best preserved in canal walls (Williams et al. 2004).
Terebratulina: Parkinson et al. (2005).
Tomteluva perturbata: No original structural details evident (Streng et al., 2016).

4.18 Sclerites: Structure

[177] Stratiform lamellae expressed at surface

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[?]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[-]Paramicrocornus[?]Haplophrentis carinatus[-]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[1]Namacalathus[-]Phoronis[2]Dailyatia[?]Cotyledion tylodes[2]Eccentrotheca[?]Orthrozanclus[1]Micromitra[2]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[-]Yuganotheca elegans[?]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[-]Orthis[?]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[?]Ussunia[?]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[1]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 177: Sclerites: Structure: Stratiform lamellae expressed at surface

1: Lamellae not expressed at surface2: Lamellae correspond to external shell ornament?: Ambiguous-: InapplicableTransformational character.

In tommotiids, the shell simply comprises a stack of stratiform lamellae, each corresponding to a circumferential rib at the shell surface. This is particularly apparent in Dailyatia (Skovsted et al., 2015) and Paterimitra (C. M. Larsson et al., 2014).

Askepasma toddense: Topper et al. (2013).
Coolinia pecten: Dewing (2004).
Dailyatia: Each lamina corresponds to a ridge on the surface (Skovsted et al., 2015).
Eoobolus: Balthasar (2007).
Novocrania: Parkinson et al. (2005).
Paterimitra: Particularly apparent (C. M. Larsson et al., 2014).
Pelagodiscus atlanticus: Williams, Cusack, & Buckman (1998a).
Salanygolina: Laminae seem to terminate at superficial ridges (Holmer et al., 2009).
Siphonobolus priscus: (Williams et al. 2004).

[178] Stratiform laminae separated

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[?]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[-]Paramicrocornus[?]Haplophrentis carinatus[-]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[1]Namacalathus[-]Phoronis[1]Dailyatia[?]Cotyledion tylodes[1]Eccentrotheca[?]Orthrozanclus[?]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[-]Yuganotheca elegans[?]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[-]Orthis[?]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[?]Ussunia[?]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 178: Sclerites: Structure: Stratiform laminae separated

1: Contiguous stratified layer2: Laminae separated by organic layers or voids-: InapplicableTransformational character.

Laminae within, for example, Salanygolina are separated by voids that may originally have contained organic material (e.g. Holmer et al., 2009). In contrast, tommotiids and paterinids exhibit stratification without voids, perhaps representing periodic fluctuations in phosphate availability (Balthasar et al., 2009).

Askepasma toddense, Eccentrotheca: Contiguous (Balthasar et al., 2009).
Botsfordia: Skovsted & Holmer 2003.
Clupeafumosus socialis: Acrotretid laminae are separated by column-supported voids.
Coolinia pecten: Dewing (2004).
Craniops: (Williams et al., 1997, fig. 249.1).
Dailyatia: Contiguous (Skovsted et al., 2015, figs. 54–55).
Eoobolus: Essentially contiguous, notwithstanding occasional laminae of calcite/chlorite (Balthasar, 2007).
Gasconsia: Hanken & Harper (1985).
Mickwitzia muralensis: (Balthasar, 2004).
Micrina: Matrix filled chambers (Balthasar et al., 2009, fig. DR1).
Mummpikia nuda: Seemingly contiguous (Balthasar, 2008).
Novocrania: Not in Novocrania, though possibly present in Neoancistrocrania (Parkinson et al., 2005).
Paterimitra: Contiguous in some regions, but large internal cavities present [Balthasar et al. (2009); see fig. DR3].
Pelagodiscus atlanticus: Williams et al. (1998a).
Salanygolina: Polygonally-filled voids (Holmer et al., 2009).
Siphonobolus priscus: (Williams et al. 2004).

[179] Stratiform laminae with polygonal ornament

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[-]Tonicella[?]Mopalia[?]Polysacos vickersianum[-]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[-]Paramicrocornus[?]Haplophrentis carinatus[-]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[1]Namacalathus[-]Phoronis[1]Dailyatia[?]Cotyledion tylodes[2]Eccentrotheca[?]Orthrozanclus[1]Micromitra[2]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[2]Salanygolina[-]Terebratulina[-]Orthis[?]Alisina[?]Longtancunella chengjiangensis[-]Glyptoria[-]Nisusia sulcata[-]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[?]Ussunia[?]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[12]Mummpikia nuda[1]Botsfordia[1]Eoobolus[?]Lingulellotreta malongensis[12]Lingula[1]Siphonobolus priscus[-]Acanthotretella spinosa[-]Lingulosacculus[2]Clupeafumosus socialis[2]Micrina[2]Paterimitra[-]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 179: Sclerites: Structure: Stratiform laminae with polygonal ornament

1: Absent2: Present?: Ambiguous-: InapplicableTransformational character.

See character 37 in Williams et al. (1998b).
“A distinct primary layer […] is characterized by a polygonal ornament that is mineralized from the polygon walls inward, while the rest of the shell and/or sclerite is secreted by basal accretion” – Balthasar et al. (2009). Distinguished from epithelial cell moulds in lingulids, which do not form an integral part of the shell structure (Balthasar et al., 2009).
Treated as transformational as ancestral condition is ambiguous.

Askepasma toddense, Eccentrotheca, Paterimitra: Present (Balthasar et al., 2009).
Micromitra, Botsfordia, Eoobolus: Lamination present, with no imprints of presumed mantle cells (following Williams et al., 1998b, appendix 2).
Clupeafumosus socialis: Epithelial cell moulds present on inner shell layer in acrotretids (Z.-L. Zhang, Zhang, & Wang, 2016).
Coolinia pecten: Dewing (2004).
Craniops: (Williams et al., 1997, fig. 249.1).
Dailyatia: Polygonal structures on external surface of sclerites only (Skovsted et al., 2015); not reported from other camenellans (Balthasar et al., 2009).
Gasconsia: Hanken & Harper (1985).
Lingula: Absent in Lingula, though potentially equivalent, if superficial (Balthasar et al., 2009), features adorn Lingulella (Curry & Williams, 1983).
Micrina: Micrina exhibits polygonal imprints on the internal surfaces of successive second-order laminae, suggesting the existence of a polygonal organization of these layers (Balthasar et al., 2009).
Mummpikia nuda: It is conceivable that the rods (Balthasar 2008) correspond to the polygonal ornament observed in other taxa; coded as ambiguous.
Novocrania: Parkinson et al. (2005).
Pelagodiscus atlanticus: Williams et al. (1998a).
Salanygolina: Prominently present (Holmer et al., 2009).
Siphonobolus priscus: (Williams et al. 2004).

[180] Canals

Character adds 6 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[1]Paramicrocornus[?]Haplophrentis carinatus[1]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[1]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[0]Eccentrotheca[?]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[?]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[?]Ussunia[?]Heliomedusa orienta[0]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[0]Botsfordia[0]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 180: Sclerites: Structure: Canals

0: Absent1: Present?: AmbiguousNeomorphic character.

A caniculate microstructure occurs in lingulids; canals are narrower (< 1 μm) than punctae, may branch, and do not fully penetrate the shell, terminating just within the boundaries of a microstructural layer. See Williams 1997, p303ff, and Balthasar 2008, p273, for discussion.

Tubules described in hyoliths by Kouchinsky (2000) measure around 10 μm in diameter, making them an order of magnitude wider than lingulid canals.

This said, Balthasar (2008) considers the rod-like tubules within the columnar shell microstructure of Mickwitzia cf. occidens (1–3 μm wide, Skovsted & Holmer 2003), acrotretides (1 μm wide, see Holmer 1989, Zhang et al. 2016) and lingulellotretids (100 nm wide, Cusack et al 1999) as equivalent to lingulid canals.

Micrina exhibits both punctae and canals (Harper et al. 2017), challenging Carlson’s contention (in Williams et al. 2007) that the structures are potentially homologous as shell perforations.

Bactrotheca: Taxon known only from moulds (Valent et al., 2012).
Botsfordia: Not evident in section presented by Skovsted & Holmer (2003).

Clupeafumosus socialis: Acrotretid laminae bear characteristic columns (e.g. Zhang et al. 2016).

Balthasar (2008) considers these columns as homologous with tubules within the columnar shell microstructure Mummpikia, Mickwitzia and lingulellotretids.
Cupitheca holocyclata: Orthogonal tubules (Vendrasco et al., 2017).
Halkieria evangelista: The chambers in halkieriid sclerites do not correspond in morphology or dimension to the brachiopod-like canals documented by this character.
Longtancunella chengjiangensis: Preservational resolution not sufficient to evaluate.
Mickwitzia muralensis: Coded as present to reflect similarity of columnar microstructure remarked on by, among others, Balthasar (2008); Williams et al. (2007); Skovsted & Holmer (2003).

Micrina: Acrotretid laminae bear characteristic columns (e.g. Zhang et al. 2016); a similar fabric has been reported, and assumed homologous, in Micrina (Butler et al. 2012).

A similar columnar shell microstructure also occurs in the closely related Mickwitzia (Balthasar 2008).
Mytilus: Gao et al. (2015).
Namacalathus: Canal-like structures have been reported in Namacalathus (Zhuravlev et al. 2015), and interpreted as evidence for a Lophophorate affinity. Though the structures are not necessarily directly equivalent, the hypothesis of homology is followed here.
Neopilina: McLean (1979).
Paramicrocornus: Columns, replicated in phosphate and present in both layers of the shell, have been interpreted as potential homologues to acrotretid columns (Z.-L. Zhang et al., 2018).
Siphonobolus priscus: The ‘canals’ through the shell have a diameter of c. 20 μm (Williams et al. 2004, text-fig. 2a), falling within the definition of punctae (rather than canals) used herein.
Tonicella: Aesthete canals do not fall within the definition of this character.

[181] Punctae

Character adds 7 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[0]Paramicrocornus[?]Haplophrentis carinatus[0]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[0]Eccentrotheca[?]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[1]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[1]Novocrania[0]Craniops[?]Ussunia[1]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[1]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[1]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 181: Sclerites: Structure: Punctae

0: Absent1: Present?: AmbiguousNeomorphic character.

Punctae are 10–20 μm wide canals created by multicellular extensions of the outer epithelium. They penetrate the full depth of the shell.

Balthasar (2008) writes:

“Vertical shell penetrating structures, such as punctae, pseudopunctae, extropunctae and canals, are common in many groups of brachiopods and are distinguished based on their geometry and size (Williams 1997). Punctae are 10–20 μm wide and represent multicellular extensions of the outer epithelium (Owen and Williams 1969). Pseudopunctae and extropunctae are similar in diameter but, instead of canals, are vertical stacks of conical deflections of individual shell layers (Williams and Brunton 1993). None of these three types of vertical shell structure, all of which are confined to calcitic-shelled brachiopods, compares with the much smaller canals (< 1 μm in diameter) of M. nuda. The only type of vertical structure that fits the size and nature of the canals of the Mural obolellids are the canals of linguliform brachiopods, which range in width from 180 to 740 nm and are occupied by proteinaceous strands in extant taxa (Williams et al. 1992; Williams et al. 1994; Williams et al. 1997). In contrast to obolellid canals, however, linguliform canals are not known to penetrate the entire shell but terminate in organic-rich layers (Williams 1997). Based on these considerations it would, therefore, be misleading to call obolellid shells punctate (they are as much”punctate" as acrotretids or other linguliforms); rather their shell structure should be called canaliculate (Williams 1997)."

Bactrotheca: Taxon known only from moulds (Valent et al., 2012).
Craniops: “impunctate”.
Haplophrentis carinatus: The tubules within the centre of the bundles of hyolith shells (Kouchinsky 2000) are c. 10 μm wide, making them an order of magnitude larger than the canals that characterize lingulid valves, and a similar scale to punctae. This said, they have only been reported in a putative allathecid, so the presence of equivalent structures in hyolithids has never been demonstrated.
Heliomedusa orienta: ‘Identical’ to those in Mickwitzia – see Williams et al. 2007.
Mickwitzia muralensis: Coded as present to reflect that the chambers contained setae; following Carlson in Williams et al. 2007, the punctae may or may not be homologous as punctae, but are likely homologous as shell perforations; both these perforations and those of Micrina were associated with setae, even if their equivalence bay be with juvenile vs adult setal structures in modern brachiopods (Balthasar 2004, p. 397).
Mummpikia nuda: “Vertical shell penetrating structures, such as punctae, pseudopunctae, extropunctae and canals, are common in many groups of brachiopods and are distinguished based on their geometry and size (Williams 1997). Punctae are 10–20 μm wide and represent multicellular extensions of the outer epithelium (Owen and Williams 1969). […] None of these three types of vertical shell structure, all of which are confined to calcitic-shelled brachiopods, compares with the much smaller canals (< 1 μm in diameter) of M. nuda. The only type of vertical structure that fits the size and nature of the canals of the Mural obolellids are the canals of linguliform brachiopods, which range in width from 180 to 740 nm and are occupied by proteinaceous strands in extant taxa (Williams et al. 1992, 1994; Williams 1997).” – Balthasar 2008.
Mytilus: Gao et al. (2015).
Neopilina: McLean (1979).
Paramicrocornus: Not evident despite excellent preservation; interpreted as absent (Z.-L. Zhang et al., 2018).
Siphonobolus priscus: The ‘canals’ through the shell have a diameter of c. 20 μm (Williams et al. 2004, text-fig. 2a), falling within the definition of punctae used herein.
Terebratulina: Endopunctae are relatively large canals, diameter vary greatly from 5–20 μm.

[182] Pseudopunctae

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[0]Paramicrocornus[?]Haplophrentis carinatus[0]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[0]Eccentrotheca[?]Orthrozanclus[0]Micromitra[0]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[?]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[?]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 182: Sclerites: Structure: Pseudopunctae

0: Absent1: PresentNeomorphic character.

Pseudopunctae are not punctae, but deflections of shell laminae. They characterise Strophomenata in particular.

Antigonambonites planus, Glyptoria, Nisusia sulcata: Scored absent in data matrix of Benedetto (2009).
Bactrotheca: Taxon known only from moulds (Valent et al., 2012).
Mytilus: Gao et al. (2015).
Neopilina: McLean (1979).
Orthis: Scored absent (in Eoorthis) in data matrix of Benedetto (2009).
Paramicrocornus: Absent (Z.-L. Zhang et al., 2018).

[183] External polygonal ornament

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[1]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 183: Sclerites: Structure: External polygonal ornament

0: Absent1: PresentNeomorphic character.

Regular polygonal compartments, around 10 μm in diameter, characterise Paterimitra. Walls between compartments have the cross-section of an anvil. An external polygonal structure (possible imprints of epithelial tissue) occurs in Dailyatia, but it is a surface pattern, which is different from the polygonal prisms in the body wall of other paterinid-like groups.

Clupeafumosus socialis: The polygonal ornament reported in acrotretids by Zhang et al. (2016) is on the internal surface of the shell.
Mytilus: Gao et al. (2015).
Neopilina: McLean (1979).

[184] Aesthete canals

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[?]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 184: Sclerites: Structure: Aesthete canals

0: Absent1: PresentNeomorphic character.

Following character 20 of Vinther et al. (2017).

Calvapilosa kroegeri: Per Vinther et al. (2017).
Halkieria evangelista: Absent in primary valves; no evidence in accessory sclerites in this taxon.
Mytilus: Gao et al. (2015).
Polysacos vickersianum: Present in spines only.
Siphogonuchites multa: Coded present by Vinther et al. (2017), who cite Bengtson (1992).
The incorporated sclerites conceivably correspond to aesthete precursors, but this cannot be decisively established: so coded as ambiguous.

[185] Aesthete canals: Orientation

Character adds 0 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[2]Tonicella[2]Mopalia[2]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[2]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-1]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 185: Sclerites: Structure: Aesthete canals: Orientation

2: Predominantly parallel to valve surface-: InapplicableTransformational character.

Following Hoare (2009).

Calvapilosa kroegeri: Seemingly parallel to valve surface (Vinther et al., 2017).
Polysacos vickersianum: Following Diadeloplax; see (???).
Siphogonuchites multa: If the spines correspond to aesthetes, they are perpendicular to the valve surface. Inapplicable otherwise.

[186] Aesthete canals: Size variation

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[2]Mopalia[1]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-2]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 186: Sclerites: Structure: Aesthete canals: Size variation

1: Uniform2: Heterogeneous?: Ambiguous-: InapplicableTransformational character.

Following Vendrasco & Runnegar (2004).

Calvapilosa kroegeri: No variation in size evident (Vinther et al., 2017).
Mopalia: Non-uniform width (Vendrasco, Fernandez, Eernisse, & Runnegar, 2008).
Tonicella: Consistently uniform (Vendrasco et al., 2008).

[187] Aesthete canals: Megalaesthete bulbs

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[-]Serpula[-]Capitella[-]Canadia spinosa[-]Sipunculus[-]Wirenia[-]Chaetoderma[1]Tonicella[1]Mopalia[2]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[-]Phthipodochiton thraivensis[-]Wiwaxia corrugata[-]Odontogriphus omalus[-]Neopilina[-]Conocardium elongatum[-]Haliotis[-]Pojetaia runnegari[-]Mytilus[-]Dentalium[-]Pelagiella[-1]Siphogonuchites multa[-]Acaenoplax hayae[-]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[-]Halkieria evangelista[-]Flustra[-]Amathia[-]Loxosomella[-]Namacalathus[-]Phoronis[-]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[-]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[-]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 187: Sclerites: Structure: Aesthete canals: Megalaesthete bulbs

1: Irregularly spaced2: Linear arrangement with orderly spacing-: InapplicableTransformational character.

Megalaesthetes are the large aesthete canals from which smaller chambers emerge. Character ‘lin’ in Vendrasco et al. (2008).

[188] Subapical tunnels

Character adds 0 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[0]Polysacos vickersianum[0]Leptochiton[0]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[0]Acaenoplax hayae[0]Kulindroplax perissokomos[0]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[0]Bactrotheca[0]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[0]Cotyledion tylodes[0]Eccentrotheca[0]Orthrozanclus[0]Micromitra[0]Askepasma toddense[0]Coolinia pecten[0]Antigonambonites planus[0]Yuganotheca elegans[0]Tomteluva perturbata[0]Salanygolina[0]Terebratulina[0]Orthis[0]Alisina[0]Longtancunella chengjiangensis[0]Glyptoria[0]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[0]Craniops[0]Ussunia[0]Heliomedusa orienta[0]Gasconsia[0]Pelagodiscus atlanticus[0]Mickwitzia muralensis[0]Mummpikia nuda[0]Botsfordia[0]Eoobolus[0]Lingulellotreta malongensis[0]Lingula[0]Siphonobolus priscus[0]Acanthotretella spinosa[0]Lingulosacculus[0]Clupeafumosus socialis[0]Micrina[0]Paterimitra[0]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 188: Sclerites: Structure: Subapical tunnels

0: AbsentNeomorphic character.

Character 23 in Vinther et al. (2017). Distinct from the umbonal perforation observed in some ventral valves on account of their subapical position. Also termed ‘lacunae’.

[189] Articulamentum

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[0]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[0]Calvapilosa kroegeri[0]Phthipodochiton thraivensis[0]Wiwaxia corrugata[0]Odontogriphus omalus[0]Neopilina[0]Conocardium elongatum[0]Haliotis[0]Pojetaia runnegari[0]Mytilus[0]Dentalium[0]Pelagiella[0]Siphogonuchites multa[?]Acaenoplax hayae[0]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[0]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[0]Namacalathus[0]Phoronis[0]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[0]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[0]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 189: Sclerites: Structure: Articulamentum

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 30 in Vinther et al. (2017). The articulamentum is a secondary layer of shell present in polyplacophorans.

Calvapilosa kroegeri: Coded as absent by Vinther et al. (2017).
Glaphurochiton carbonarius, Polysacos vickersianum: Following Vinther et al. (2017).
Kulindroplax perissokomos: Sutton et al. (2012).
Phthipodochiton thraivensis: Absent (Sutton & Sigwart, 2012).

4.19 Gametes

[190] Gonocoel

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 190: Gametes: Gonocoel

0: Absent1: Retroperineal gonads?: AmbiguousNeomorphic character.

Character 27 in Haszprunar (1996).

[191] Gonad position

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[?]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[1]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 191: Gametes: Gonad position

1: Dorsal to gut2: Ventral to gut?: AmbiguousTransformational character.

Character 32 in Haszprunar (2000).

[192] Ovary wall saccular

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[1]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 192: Gametes: Ovary wall saccular

0: Plain1: Saccular?: AmbiguousNeomorphic character.

After character 31 in Haszprunar (1996).

[193] Testis wall saccular

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[1]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 193: Gametes: Testis wall saccular

0: Plain1: Saccular?: AmbiguousNeomorphic character.

After character 31 in Haszprunar (1996).

[194] Asexual reproduction

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[0]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[0]Haplophrentis carinatus[0]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[0]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[0]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 194: Gametes: Asexual reproduction

0: Never exhibited1: Frequently exhibited?: AmbiguousNeomorphic character.

After character 30 in Haszprunar (1996).

Namacalathus: Budding well documented (e.g. Zhuravlev, Wood, & Penny, 2015).

[195] Sexes

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[2]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 195: Gametes: Sexes

1: Gonochoristic2: Hermaphroditic?: AmbiguousTransformational character.

After characters 1.61 and 2.54 in von Salvini-Plawen & Steiner (1996).

Amathia: Hermaphroditic (Reed, 1988).

[196] Fertilization

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[2]Capitella[?]Canadia spinosa[1]Sipunculus[2]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 196: Gametes: Fertilization

1: External2: Internal?: AmbiguousTransformational character.

After character 62 in Haszprunar (2000).

Amathia: Brood pouches in abandoned lophophore.

4.20 Gametes: Egg

[197] Size

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[2]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[2]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[1]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 197: Gametes: Egg: Size

1: Small: < 100 um, little yolk2: Large: > 110 um, much yolk?: AmbiguousTransformational character.

Following Carlson (1995), character 7. This character is only possible to code in extant taxa. It is not considered independent of Carlson’s character 11, number of gametes released per spawning, as it is possible to produce more small eggs than large eggs – thus this latter character is not reproduced in the present study. The same goes for Carlson’s character 12, gamete dispersal mode; brooders will tend to brood large eggs.

Amathia: “Mature eggs commonly measure about 200 μm in diameter” (Franzén, 1977); the larva is a similar size (Reed & Cloney, 1982).
Capitella: >200 μm in most species (though 50 μm in some) (Eckelbarger & Grassle, 1983).
Dentalium: Egg size can vary from 60–200 μm in scaphopods, but in Dentalium the eggs are large (Dufresne-Dube, Picheral, & Guerrier, 1983).
Flustra: “Mature eggs commonly measure about 200 μm in diameter” – Franzén (1977).
Haliotis: Up to 200 μm long when fully developed (Martin, Romero, & Miller-Walker, 1983).
Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina: Following coding for class in Carlson (1995) appendix 1, character 7.
Loxosomella: Tiny (Nielsen, 1966).
Mytilus: c. 30 μm (Humphreys, 1962).
Neopilina: “Usually the mature eggs have an oblong cell body 220–320 μm long and 130–190 μm broad.” (Lemche & Wingstrand, 1959).
Phoronis: Phoronis has planktotrophic larvae. indicating a small egg size (Ruppert et al. 2004). Carlson (1995) codes phoronids as polymorphic, as some members of the phylum have eggs of each size.
Serpula: c. 50 μm in Hydroides (Miles, Hadfield, & Wayne, 2007).
Siphonobolus priscus: “the ventral brephic valve [was] 50 μm across, [which] is close to the known lower limit of the brachiopod egg size” – Popov et al. 2009.
Sipunculus: c. 200 μm in diameter (Rice, 1988).
Tonicella: Buckland‐Nicks, Koss, & Chia (1988).

[198] Protective membrane

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 198: Gametes: Egg: Protective membrane

0: Absent1: Present?: AmbiguousNeomorphic character.

After character 4.69 in von Salvini-Plawen & Steiner (1996).

Flustra, Amathia: “Eggs have a loose consistency and are capable of changing form” (Franzén, 1977).
Capitella: Eckelbarger & Grassle (1983).
Haliotis: Surrounded by external gelatinous envelope, whose homology to the protective membrane of other taxa is uncertain – hence coded ambiguous.
Phoronis: Eggs “are surrounded by a delicate fertilization membrane” (Pennerstorfer & Scholtz, 2012).

[199] Site of maturation

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[?]Sipunculus[?]Wirenia[?]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[4]Neopilina[?]Conocardium elongatum[4]Haliotis[?]Pojetaia runnegari[4]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[3]Flustra[3]Amathia[?]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 199: Gametes: Egg: Site of maturation

1: Body cavity2: Mantle canals3: Ovicell4: Ovaries?: AmbiguousTransformational character.

After Carlson (1995), character 9. Only possible to code in extant taxa.

Flustra, Amathia: Ovicell (Franzén, 1977).
Capitella: “Most ultrastructural features of the eggs in the lateral region of the ovary are indistinguishable from those floating freely in the coelom, although the egg envelopes […] undergo additional differentiation following ovulation […] there is no indication that further maturation occurs before spawning” (Eckelbarger & Grassle, 1983).
Haliotis: Compartments of the ovary wall (Martin et al., 1983).
Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina: Following Hodgson & Reunov (1994).
Mytilus: Mature eggs within the ovary (Humphreys, 1962).
Neopilina: Lemche & Wingstrand (1959).
Phoronis: Following coding for class in Carlson (1995) Appendix 1, character 9.

[200] Nucleus: Aspect

Character adds 6 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 200: Gametes: Spermatozoa: Nucleus: Aspect

0: Equant: length comparable to width1: Elongate: length more than twice width?: AmbiguousNeomorphic character.

After character 41 in Ponder & Lindberg (1997).

Flustra, Amathia: Elongate (Franzén, 1981).
Capitella: Eckelbarger & Grassle (1987).
Dentalium: Elongate nucleus, 4–6 times longer than wide (Dufresne-Dube et al., 1983).
Haliotis: At least three times longer than wide in the gastropods figured in Healy, Schaefer, & Haszprunar (1995).
Loxosomella: Elongate in Loxosoma (Franzén, 2000).
Mytilus: Around five times longer than wide in Nucula (Healy et al., 1995).
Neopilina: Only a little longer than wide in Laevipilina (Healy et al., 1995).
Serpula: Gherardi et al. (2011).
Tonicella: Profoundly elongated nucleus (Buckland‐Nicks et al., 1988).

4.21 Gametes: Spermatozoa

[201] Nucleus: Shape

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[2]Capitella[?]Canadia spinosa[?]Sipunculus[2]Wirenia[2]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[?]Amathia[2]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 201: Gametes: Spermatozoa: Nucleus: Shape

1: Convex (i.e. round)2: Concave (i.e. with invagination)?: AmbiguousTransformational character.

Most spermatozoa have nuclei with an invagination; see character 50 in Ponder & Lindberg (1997).

Capitella: Eckelbarger & Grassle (1987).
Dentalium: Healy et al. (1995).
Haliotis: Following gastropods illustrated in Healy et al. (1995).
Neopilina: Following Laevipilina (Healy et al., 1995).
Tonicella: Buckland‐Nicks et al. (1988).

[202] Nucleus: Nuclear filament

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[?]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 202: Gametes: Spermatozoa: Nucleus: Nuclear filament

0: Absent1: PresentNeomorphic character.

A nuclear filament is an anterior extension of the nucleus that terminates at the acrosome, present in lepidopleurid chitons (Buckland-Nicks, 2008, character 6).

Capitella: Eckelbarger & Grassle (1987).
Wirenia, Chaetoderma: Buckland-Nicks (2008).
Haliotis: Lewis, Leighton, & Vacquier (1980).
Leptochiton: Absent (Buckland-Nicks, 2008, fig. 2A).
Mopalia: Prominent (Buckland-Nicks, 2008, fig. 3D).
Mytilus: Niijima (1965).
Tonicella: Present (Buckland-Nicks, 2008).

[203] Anterior nuclear fossa

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[1]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 203: Gametes: Spermatozoa: Anterior nuclear fossa

0: Absent1: Present?: AmbiguousNeomorphic character.

After character 160 in Giribet & Wheeler (2002). A fossa (latin: ditch) is a dent or impression.

Flustra, Amathia: Present (in Tubulipora; Franzén, 1984).
Dentalium: Dufresne-Dube et al. (1983).
Haliotis: Lewis et al. (1980).
Loxosomella: Present in Loxosoma (Franzén, 2000).
Mytilus: Deep “tubular passage through centre of nucleus” (Niijima, 1965).
Neopilina: Following Laevipilina in Healy et al. (1995).
Pelagodiscus atlanticus: Present in Discinisca tenuis (Hodgson & Reunov, 1994).
Phoronis: Nucleus “almost round” (Reunov & Klepal, 2004).
Serpula: Absent: subacrosomal space does not impinge on nuclear envelope (Gherardi et al., 2011).
Sipunculus: Prominent in Phascolion (Rice, 1993).
Terebratulina: No anterior invagination (Hodgson & Reunov, 1994).
Tonicella: Buckland‐Nicks et al. (1988).

[204] Acrosome

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 204: Gametes: Spermatozoa: Acrosome

0: Absent1: PresentNeomorphic character.

Sometimes fully termed the Acrosome vesicle.

Wirenia, Chaetoderma, Mopalia, Tonicella, Leptochiton: Buckland-Nicks (2008), character 1.

[205] Acrosome: Shape

Character adds 8 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[3]Serpula[4]Capitella[?]Canadia spinosa[3]Sipunculus[4]Wirenia[5]Chaetoderma[5]Tonicella[5]Mopalia[?]Polysacos vickersianum[4]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[4]Neopilina[?]Conocardium elongatum[4]Haliotis[?]Pojetaia runnegari[4]Mytilus[4]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[4]Flustra[4]Amathia[4]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[3]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 205: Gametes: Spermatozoa: Acrosome: Shape

1: Pear-shaped2: Needle-shaped3: Disc-shaped4: Conical5: Vesicle, i.e. subspherical?: AmbiguousTransformational character.

Flustra, Amathia: Conical (in Tubulipora; Franzén, 1984).
Capitella: Eckelbarger & Grassle (1987).
Chaetoderma, Mopalia, Tonicella: Vesicular (Buckland-Nicks, 2008, character 2).
Dentalium: Low conical aspect (Dufresne-Dube et al., 1983).
Haliotis: Conical in gastropods (Healy et al., 1995).
Leptochiton: Conical (Buckland-Nicks, 2008, character 2).
Lingula: Pear-shaped (Fukumoto 2003).
Loxosomella: Conical/cylindrical acrosome-like extension in Loxosoma (Franzén, 2000).
Mytilus: Conical in Nucula (Healy et al., 1995).
Neopilina: Conical in Laevipilina (Healy et al., 1995).
Novocrania: Needle-shaped (Afzelius & Ferraguti, 1978).
Pelagodiscus atlanticus: Pear-shaped (Hodgson & Reunov, 1994).
Phoronis: Needle-shaped (Reunov & Klepal, 2004).
Serpula: Gherardi et al. (2011).
Sipunculus: A peaked disc in Phascolion (Rice, 1993).
Terebratulina: Disc-shaped (in Kraussina) (Hodgson & Reunov, 1994).
Wirenia: Conical (in Epimenia; Buckland-Nicks, 2008, character 2).

[206] Acrosome: Differentiated internally

Character adds 7 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 206: Gametes: Spermatozoa: Acrosome: Differentiated internally

0: No internal differentiation1: Acrosome differentiated internally?: AmbiguousNeomorphic character.

Hodgson & Reunov 1994 describe the Discinisca acrosome as having “an electron-lucent centre and an electron-dense outer region”, and state that this trait is characteristic of inarticulate brachiopods. The interstitial granule of certain polyplacophorans represents a separate mode of acrosome differentiation. The subacrosomal granule and subacrosomal basal plate are treated separately, and are not considered to represent internal differentiation.

Flustra, Amathia: No evidence of internal differentiation (in Tubulipora; Franzén, 1984).
Capitella: Electron dense rings the the acrosome vesicle (Eckelbarger & Grassle, 1987).
Chaetoderma: Not differentiated, following character 2 of Buckland-Nicks (2008).
Dentalium: Differentiated membrane only (Dufresne-Dube et al., 1983).
Haliotis: “The large acrosome granule contains two distinct components: (1) an ovoid electron-dense body in the anterior apex of the granule […], and (2) a less dense, homogeneous material at the granule posterior.” (Lewis et al., 1980).
Leptochiton: The acrosome is a cone with subacrosomal granule and subacrosomal plate, but not interstital granule, following character 2 of Buckland-Nicks (2008).
Lingula: Clear differentiation of marginal area (Fukumoto 2003).
Loxosomella: Not evident in Loxosoma (Franzén, 2000).
Mopalia: The acrosome is a cone with subacrosomal granule, interstitial granule, and subacrosomal plate, following character 2 of Buckland-Nicks (2008).
Mytilus: Material lines acrosomal membrane (Niijima, 1965).
Neopilina: Not consistently differentiated in Laevipilina (Healy et al., 1995).
Novocrania: “Along the inner and outer margins there are periodically banded layers, and between them there is a less dense layer” – Afzelius & Ferraguti, 1978.
Pelagodiscus atlanticus: Following Discinisca tenuis, described in Hodgson & Reunov (1994).
Phoronis: Acrosome-like structure; no internal division or surrounding membrane, with possibility that much of the acrosome is secondarily lost (Reunov & Klepal 2004).
Serpula: Gherardi et al. (2011).
Sipunculus: No differentiation within acrosomal vesicle (Rice, 1993).
Terebratulina: Following Hodgson & Reunov (1994).
Tonicella: “One can distinguish two components in the acrosome, an apical and a basal granule” – Buckland‐Nicks et al. (1988).
Wirenia: In Epimenia, the acrosome is a cone with subacrosomal granule, interstitial granule, and subacrosomal plate, following character 2 of Buckland-Nicks (2008).

[207] Acrosome: Subacrosomal basal plate

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[?]Sipunculus[1]Wirenia[1]Chaetoderma[0]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[1]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 207: Gametes: Spermatozoa: Acrosome: Subacrosomal basal plate

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 41 in Ponder & Lindberg (1997).

Capitella: Eckelbarger & Grassle (1987).
Haliotis: Axial filament but not basal plate (Lewis et al., 1980).
Mytilus: Basal ring (Niijima, 1965).
Neopilina: Coded following Laevipilina (Healy et al., 1995).

[208] Acrosome: Subacrosomal basal plate: Basal granule

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[?]Sipunculus[1]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 208: Gametes: Spermatozoa: Acrosome: Subacrosomal basal plate: Basal granule

0: Absent1: PresentNeomorphic character.

In certain taxa, the subacrosomal basal plate develops a subacrosomal granule (Buckland-Nicks, 2008).

[209] Mid-piece

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 209: Gametes: Spermatozoa: Mid-piece

0: Multiple mitochondria1: Single ring-shaped mitochondrion?: AmbiguousNeomorphic character.

Following Hodgson & Reunov (1994).

Flustra, Amathia: Two mitochondrial derivatives in Flustra (Franzén, 1977, 1981); four in Tubulipora (Franzén, 1984).
Capitella: Single ring-shaped mitochondrion (Eckelbarger & Grassle, 1987).
Chaetoderma, Mopalia, Tonicella, Leptochiton: Buckland-Nicks (2008).
Dentalium: Dufresne-Dube et al. (1983).
Haliotis: Five mitochondria (Lewis et al., 1980).
Lingula, Terebratulina: Following Hodgson & Reunov (1994).
Loxosomella: “The midpiece consists of two long mitochondrial rods connected with each other by a thin mitochondrial lamella” (Franzén, 2000, in Loxosoma); these are essentially a single organelle surrounding a central rod of electron-dense material.
Mytilus: Five mitochondria (Niijima, 1965).
Neopilina: Five mitochondria in Laevipilina (Healy et al., 1995).
Novocrania: Four mitochondria (Afzelius & Ferraguti, 1978).
Pelagodiscus atlanticus: Following Discinisca tenuis, described in Hodgson & Reunov (1994).
Phoronis: The mitochondria fuse in the middle stage of spermiogenesis to become a pair of mitochondria (Reunov & Klepal 2004).
Serpula: Five mitochondria in ring (Gherardi et al., 2011).
Sipunculus: Ring of five mitochondria around the central centriole (Rice, 1993).

[210] Mid-piece: Mitochondrial location

Character adds 5 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[?]Sipunculus[1]Wirenia[2]Chaetoderma[4]Tonicella[4]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[2]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[?]Amathia[3]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 210: Gametes: Spermatozoa: Mid-piece: Mitochondrial location

1: In sheath around flagellum2: In ring around centrioles3: In ring around offset basal body4: Lateral and anterior mitochondriaTransformational character.

Following character 3 in Buckland-Nicks (2008) and character 166 in Giribet & Wheeler (2002).

Capitella: Eckelbarger & Grassle (1987).
Haliotis: (Lewis et al., 1980).
Mytilus: Niijima (1965).
Neopilina: Coded following Laevipilina (Healy et al., 1995).
Tonicella: (Buckland-Nicks, 2008).

[211] Centrioles: Orientation

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[0]Capitella[?]Canadia spinosa[?]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 211: Gametes: Spermatozoa: Centrioles: Orientation

0: Orthogonal1: Parallel?: AmbiguousNeomorphic character.

Following Hodgson & Reunov 1994.

Flustra, Amathia: (Franzén, 1981).
Capitella: At a 90° angle (Eckelbarger & Grassle, 1987).
Dentalium: Dufresne-Dube et al. (1983).
Haliotis: Orthogonal (Lewis et al., 1980).
Leptochiton: Buckland-Nicks (2008), fig. 1B.
Lingula, Terebratulina: Following Hodgson & Reunov (1994).
Mopalia: Buckland-Nicks (2008), fig. 1D.
Mytilus: Niijima (1965).
Neopilina: Coded following Laevipilina (Healy et al., 1995).
Novocrania: Two orthogonal centrioles (Afzelius & Ferraguti 1978).
Pelagodiscus atlanticus: Following Discinisca tenuis, described in Hodgson & Reunov (1994).
Phoronis: Only one centriole in spermatzoon (Reunov & Klepal 2004, p. 7), but centrioles are perpendicularly oriented in spermatogonia (ibid., p. 2).
Serpula: The proximal centriole is parallel to the flagellum (Gherardi et al., 2011).
Tonicella: Buckland‐Nicks et al. (1988).

[212] Centrioles: Fusion

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[1]Sipunculus[0]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 212: Gametes: Spermatozoa: Centrioles: Fusion

0: Discrete1: FusedNeomorphic character.

Following character 9 in Buckland-Nicks (2008).

Sipunculus, Flustra, Amathia: Proximal centriole fused anterior to distal centriole.
Capitella: Eckelbarger & Grassle (1987).
Chaetoderma, Leptochiton: Separate centrioles (Buckland-Nicks, 2008).
Dentalium: Proximal centriole fused anterior to distal centriole (Dufresne-Dube et al., 1983).
Novocrania, Lingula, Phoronis, Loxosomella: Basal body in deep nuclear fossa.
Mopalia: Buckland-Nicks (2008).
Mytilus: Niijima (1965).
Neopilina: Coded following Laevipilina (Healy et al., 1995).
Tonicella: Proximal centriole fused lateral to distal centriole and offset (Buckland-Nicks, 2008).
Wirenia: Basal body in deep nuclear fossa (in Epimenia, Buckland-Nicks, 2008).

[213] Satellite fibre complex

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 213: Gametes: Spermatozoa: Satellite fibre complex

0: Annulus not associated with satellite fibres1: Annulus associated with satellite fibresNeomorphic character.

Following Smith (2012a), after character 48 in Ponder & Lindberg (1997).

Haliotis: Ten anchors or “satellite bodies” surround the distal centriole, two attaching to each mitochondrion (Lewis et al., 1980).
Neopilina: Coded following Laevipilina (Healy et al., 1995).

[214] Mitochondria: Shape

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[3]Capitella[?]Canadia spinosa[1]Sipunculus[4]Wirenia[1]Chaetoderma[3]Tonicella[3]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 214: Gametes: Spermatozoa: Mitochondria: Shape

1: Spherical to subspherical2: Rods3: Elongate, sac-like4: Fused spiral?: AmbiguousTransformational character.

After character 5 in Buckland-Nicks (2008); see also character 43 in Ponder & Lindberg (1997).

Flustra, Amathia: Rods (Franzén, 1981).
Haliotis: (Lewis et al., 1980).
Loxosomella: Elongate rods in Loxosoma (Franzén, 2000).
Mytilus: Niijima (1965).
Neopilina: Coded following Laevipilina (Healy et al., 1995).
Tonicella: See Buckland‐Nicks et al. (1988).

[215] Mitochondria: Cristae: Configuration

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 215: Gametes: Spermatozoa: Mitochondria: Cristae: Configuration

0: Unmodified1: Arranged in parallel platesNeomorphic character.

After character 44 in Ponder & Lindberg (1997). Cristae are internal compartments formed by inner mitochondrial membranes.

Flustra, Amathia: “Typical cristae”; “Randomly oriented” – Franzén (1984) (in Tubulipora).
Haliotis: Lewis et al. (1980).
Mopalia, Tonicella, Leptochiton: Buckland-Nicks (2008).
Loxosomella: in Loxosoma (Franzén, 2000).
Neopilina: Arranged in “a loose, radial pattern” in Laevipilina (Healy et al., 1995).

[216] Mitochondria: Midpiece

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[2]Capitella[?]Canadia spinosa[1]Sipunculus[3]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[?]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[3]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[3]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 216: Gametes: Spermatozoa: Mitochondria: Midpiece

1: Extremely short2: Long3: Forms continuous sheath?: AmbiguousTransformational character.

After Smith (2012a); see also character 43 in Ponder & Lindberg (1997); character 164 in Giribet & Wheeler (2002).

Flustra, Amathia: Long (Franzén, 1981).
Capitella: Long cytoplasmic collar (Eckelbarger & Grassle, 1987).
Haliotis: (Lewis et al., 1980).
Loxosomella: As long as the flagellum in Loxosoma (Franzén, 2000).
Mytilus: Niijima (1965).
Neopilina: Coded following Laevipilina (Healy et al., 1995).
Serpula: Five mitochondria surround the base of the flagellum in short midpiece, comparable to that of Sipunculus and Dentalium (Gherardi et al., 2011).
Sipunculus: Short ring of five mitochondria around the central centriole (Rice, 1993).

4.22 Embryo

[217] Micromere size

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[2]Sipunculus[1]Wirenia[?]Chaetoderma[2]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[2]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 217: Embryo: Micromere size

1: Similar to macromeres2: Small relative to macromeres?: AmbiguousTransformational character.

Following Hejnol (2010). Blastomeres may undergo significant size differentiation, generating macromeres and micromeres of prominently different sizes.

Flustra, Amathia: In Membranipora, “cleavage is slightly unequal resulting in little larger central
blastomeres” (Gruhl, 2010b).
Capitella: Meyer, Boyle, Martindale, & Seaver (2010).
Haliotis: Biggelaar (1993).
Lingula, Terebratulina: Williams et al. (1997).
Phoronis: Uniform size (Pennerstorfer & Scholtz, 2012).
Sipunculus: Prominent differentiation in Phascolosoma (Adrianov, Maiorova, & Malakhov, 2011).
Wirenia: Similar in size, with possible exception of macromere 1D, by incorporation of the polar lobe (in Epimenia (Okusu, 2002)).

[218] Equal

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[2]Sipunculus[1]Wirenia[?]Chaetoderma[2]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[?]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 218: Embryo: Cleavage: Equal

1: Unequal2: Equal?: AmbiguousTransformational character.

Following character 170 in Giribet & Wheeler (2002).

Capitella: Meyer et al. (2010).
Haliotis: “The first two cleavages are equal” (Biggelaar, 1993).
Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina: Equal, in all brachiopods (Williams et al., 1997).
Mytilus: Unequal in Acila, Nucula and Yoldia (Zardus & Morse, 1998).
Phoronis: “Cleavage is holoblastic and results in approximately equal sized, or adequal, blastomeres.” – Pennerstorfer & Scholtz (2012).
Wirenia: Unequal in Epimenia (Okusu, 2002).

[219] Cross pattern

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 219: Embryo: Cleavage: Cross pattern

0: Absent1: Cross, whether “molluscan” or “annelid”?: AmbiguousNeomorphic character.

The “molluscan cross” and “annelid cross” cannot be systematically discriminated from one another, so are treated as a single state.
See characters 127 & 128 in Rouse (1999); 1.49 in von Salvini-Plawen & Steiner (1996);
character 34 in Haszprunar (1996); 35 in Haszprunar (2000); 172 in Giribet & Wheeler (2002).

Capitella: Meyer et al. (2010).
Haliotis: “Cross-furrow” pattern (Biggelaar, 1993).
Mytilus: Absent in Acila, Nucula and Yoldia (Zardus & Morse, 1998).
Wirenia: Visible but indistinct cross-furrow in Epimenia (Okusu, 2002).

[220] Polar lobe formation

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[2]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[?]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 220: Embryo: Cleavage: Polar lobe formation

1: Absent2: Present?: AmbiguousTransformational character.

Following character 171 in Giribet & Wheeler (2002).

Haliotis: Biggelaar (1993).
Mytilus: Present in Acila, Nucula and Yoldia (Zardus & Morse, 1998).
Wirenia: Two polar lobes formed in Epimenia (Okusu, 2002).

[221] Spiral

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[2]Sipunculus[2]Wirenia[?]Chaetoderma[2]Tonicella[2]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 221: Embryo: Cleavage: Spiral

1: Absent2: Present?: AmbiguousTransformational character.

See characters 32–33 in Haszprunar (1996); character 1.48 in von Salvini-Plawen & Steiner (1996); character 29 in Glenner et al. (2004).

Flustra, Amathia: “While entoprocts are spiral cleavers, ectoprocts show a radial cleavage pattern” – Fuchs & Wanninger (2008).
Chaetoderma: Following closest relative in Glenner et al. (2004).
Phoronis: “The observed cleavage displays several characters consistent with the pattern of spiral cleavage” (Pennerstorfer & Scholtz, 2012).
Wirenia: Cleavage is spiral in Epimenia (Okusu, 2002).

[222] Origin of mesoderm

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 222: Embryo: Origin of mesoderm

1: 4d cell, from the blastopore ridge, or as ectomesoderm2: Archenteron?: AmbiguousTransformational character.

After characters 32 in Grobe (2007) and 36–37 in Glenner et al. (2004).

Pelagodiscus atlanticus, Phoronis, Sipunculus, Loxosomella, Flustra, Amathia, Mytilus, Haliotis: Following closest relative in Glenner et al. (2004).
Capitella: Four to seven origins of mesoderm, all ectomesodermal (Meyer et al., 2010).
Terebratulina: Williams et al. (1997).

4.23 Larva: Apical organ

[223] Muscles extending to the hyposphere

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 223: Larva: Apical organ: Muscles extending to the hyposphere

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 8 in Vinther et al. (2008).

Flustra, Amathia: Median muscles extending from apical organ (Gruhl, 2008).
Dentalium: Apical organ has disappeared before musculature is set in place (Wanninger & Haszprunar, 2002b).
Phoronis: Not evident (Santagata, 2004, fig. 2C).

[224] Serotonergic cells

Character adds 7 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[3]Sipunculus[?]Wirenia[?]Chaetoderma[3]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[4]Haliotis[?]Pojetaia runnegari[2]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[4]Flustra[4]Amathia[3]Loxosomella[?]Namacalathus[4]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[3]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[4]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 224: Larva: Apical organ: Serotonergic cells

1: Two flask-shaped cells2: Four flask-shaped cells3: Cluster of c. eight flask-shaped cells4: Aggregation of multiple cells of multiple types?: AmbiguousTransformational character.

Character 8 in Haszprunar & Wanninger (2008).

Flustra, Amathia: Concentration of 30–40 serotonergic perikarya (in Fredericella; Gruhl, 2010a).
Capitella: Two in Platynereis (Marlow et al., 2014).
Haliotis: “As early as Day 5 of embryogenesis, whole-mount ICC revealed an unpaired median cell (UMC) and a bilateral pair of cells that are immunoreactive for serotonin in the anterior region of the animal. Each of the three cells sends one anterior projection and one central projection that forms a dense plexus of serotonergic neurites. By Day 7, a second pair of bilateral cells is added slightly medially and posteriorly to the first pair. [… At] the last larval stage […] additional serotonergic neurons (usually two) [are] detected on each side of the five serotonergic cells.” (Marois & Carew, 1997) The cells are not obviously flask shaped.
Lingula: Cluster of “numerous” serotonergic cells (Altenburger & Wanninger, 2010; Hay-Schmidt, 1992); more than, but probably equivalent to, the flask-shaped cells of Terebratalia (Lüter, 2016).
Loxosomella: Six to eight apical cells; eight peripheral cells (Wanninger, Fuchs, & Haszprunar, 2007), indicating a probable equivalence to polyplacophorans (Haszprunar & Wanninger, 2008).
Mytilus: Four to five vase-shaped cells (Voronezhskaya, Nezlin, Odintsova, Plummer, & Croll, 2008).
Novocrania: Four flask-shaped cells (Altenburger & Wanninger, 2010).
Phoronis: Multiple shapes of cells present (Santagata, 2002); resembles the linguliform arrangement (Altenburger & Wanninger, 2010).
Sipunculus: Cluster of around eight cells, though not quite countable (Wanninger, Koop, Bromham, Noonan, & Degnan, 2005).
Terebratulina: Eight in Terebratalia (Lüter, 2016).
Tonicella: Eight in Ischnochiton and Mopalia (Wanninger et al., 2007).

[225] Develops into adult brain

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[2]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 225: Larva: Apical organ: Develops into adult brain

1: Brain has other origin2: Adult brain derived from larval apical organ / apical pole?: AmbiguousTransformational character.

Character 79 in Glenner et al. (2004).

Capitella: In Platynereis (Marlow et al., 2014).
Mytilus, Haliotis: Following closest relative in Glenner et al. (2004).
Lingula: “both the larval apical ganglion and the ventral ganglion must be retained as
the adult nervous system” (Hay-Schmidt, 1992), but not necessarily as the brain.

[226] Brain persists into adulthood

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[2]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[?]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 226: Larva: Brain persists into adulthood

1: Brain lost2: Brain retained to adulthood?: AmbiguousTransformational character.

After character 3 in Richter et al. (2010).

Haliotis: Barlow & Truman (1992).

[227] Origin of body cavity

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[1]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 227: Larva: Origin of body cavity

1: Mesenchyme2: Coelom?: AmbiguousTransformational character.

Character 1.43 in@SPS1996.

[228] Formation of coelomoducts

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[-]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 228: Larva: Formation of coelomoducts

1: Outgrowth2: Ingrowth?: Ambiguous-: InapplicableTransformational character.

Character 26 in Haszprunar (2000).

Loxosomella: Coelomoducts absent (Haszprunar, 2000).

[229] Foot

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[?]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 229: Larva: Foot

0: Absent1: Present?: AmbiguousNeomorphic character.

Foot or neurotroch present in larval stage, whether or not it is also present in mature individuals. Following Wingstrand (1985).

Capitella: The neurotroch (Meyer et al., 2010) is a cilliated ventral ‘foot’.
Loxosomella: A foot is present in the creeping-type larva of Loxosomella murmanica, though absent in L. atkinsae and the many other entoprocts that have swimming-type larvae (Fuchs & Wanninger, 2008).
Sipunculus, Serpula: Wingstrand (1985) considers the annelid neurotroch to be potentially homologous with the molluscan and entoproct foot.
Wirenia: Ciliated foot present in larvae of Epimenia (Okusu, 2002).

[230] Pedal gland

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[1]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 230: Larva: Foot: Pedal gland

0: Absent1: PresentNeomorphic character.

A pedal gland is considered evidence for homology between the molluscan and entoproct foot (Haszprunar & Wanninger, 2008).

Amathia: Ciliated clef corresponds to position of foot (Reed & Cloney, 1982), but dedicated foot not present.
Chaetoderma: Lacking (Wingstrand, 1985).
Mopalia, Tonicella, Leptochiton: “Absent in adult Polyplacophora but present in early ontogenetic stages” (Wingstrand, 1985).
Wirenia: Pedal pit present in Epimenia (Okusu, 2002).

[231] Pedal gland: Retained to adulthood

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[1]Wirenia[0]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 231: Larva: Foot: Pedal gland: Retained to adulthood

0: Absent1: PresentNeomorphic character.

Characters 1.13, 1.40 & 2.08 in A. H. Scheltema (1993); 114 in Giribet & Wheeler (2002); 1.53 in von Salvini-Plawen & Steiner (1996); 9 in Haszprunar (1996).

Chaetoderma: “Caudofoveata certainly lack […] the pedal gland” (Wingstrand, 1985).
Mopalia, Tonicella, Leptochiton: “Absent in adult Polyplacophora but present in early ontogenetic stages” (Wingstrand, 1985).
Neopilina: Lemche & Wingstrand (1959).

[232] Paired

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 232: Larva: Coelom: Paired

0: Absent1: Paired coelom originating from two teloblasts derived from 4d?: AmbiguousNeomorphic character.

Character 2.02 in A. H. Scheltema (1993).

Amathia: No evidence of pairing (Reed & Cloney, 1982).
Capitella, Mopalia, Tonicella, Leptochiton, Dentalium, Mytilus, Haliotis: Following figure 13 in A. H. Scheltema (1993).
Flustra: Hypostegal coelom separated from principal (perigastric) body cavity in cheilostomata – but this is not clearly equivalent to the paired coelom intended by this character. The coelom of Fredericella is not paired (Gruhl, 2010a).

[233] Paried: Includes pericardium

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 233: Larva: Coelom: Paried: Includes pericardium

0: Paired coelom absent, or does not include pericardium1: Paired coelom includes pericardiumNeomorphic character.

Character 1.03 in A. H. Scheltema (1993).

Leptochiton, Neopilina, Mytilus, Haliotis: Following figure 12 in A. H. Scheltema (1993).

[234] Feeding

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 234: Larva: Feeding

1: Lecithotrophic (or otherwise non-feeding)2: Planktotrophic (or otherwise feeding)?: AmbiguousTransformational character.

Character 140 in Rouse (1999). See also character 2.66 in von Salvini-Plawen & Steiner (1996); 153 in Giribet & Wheeler (2002).

Amathia: Lecithotrophic (Reed & Cloney, 1982).
Flustra: Metamorphose almost immediately after release from gonozooid (Zimmer & Woollacott, 2013); most bryozoans are lecithotrophic (Reed & Cloney, 1982).
Haliotis: (???).

4.24 Larva: Cilia

[235] Metatroch

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[0]Capitella[?]Canadia spinosa[1]Sipunculus[0]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 235: Larva: Cilia: Metatroch

0: Absent1: Present?: AmbiguousNeomorphic character.

See characters 129 and 131 in Rouse (1999); 40 in Haszprunar (1996).
A prototroch is the defining character of a trochophore larva; a metatroch is a secondary ciliary ring (Rouse, 1999).

The metatroch is present in a subset of annelids; in Polygordius, it derives from the 3c and 3d micromeres, whereas in molluscs the secondary ciliary band derives frmo 2a, 2b and 2c (Meyer et al., 2010). As such, the structures may not be homologous between molluscs and annelids.

Capitella: Absent (Marlow et al., 2014).
Terebratulina: Williams et al. (1997).
Wirenia: Prototroch only in Epimenia (Okusu, 2002).

[236] Telotroch

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[1]Wirenia[1]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 236: Larva: Cilia: Telotroch

0: Absent1: Present?: AmbiguousNeomorphic character.

A posterior ciliary band. Character 136 in Rouse (1999).

Amathia: Absent; single ciliary field lacks telotroch equivalent (Reed & Cloney, 1982).
Flustra: Absent (Zimmer & Woollacott, 2013).
Haliotis: De Viçose, Viera, Bilbao, & Izquierdo (2007).
Serpula: Following Parry & Caron (2019).
Terebratulina: Williams et al. (1997).
Wirenia: Present in Epimenia (Okusu, 2002).

[237] Ciliated food groove

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[0]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 237: Larva: Cilia: Ciliated food groove

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 132 in Rouse (1999).

Amathia: Coronal cilia do not form a food groove (Reed & Cloney, 1982).
Flustra: Cyclostomes are covered in cilia but not arranged in food groove.
Serpula: Following Parry & Caron (2019).
Terebratulina: Williams et al. (1997).
Wirenia: Not present in Epimenia (Okusu, 2002).

[238] Ciliary bands: Downstream

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 238: Larva: Cilia: Ciliary bands: Downstream

0: Absent1: Present?: AmbiguousNeomorphic character.

Downstream-collecting ciliary bands of compound cilia on multiciliated cells. Character 31 in Glenner et al. (2004).

Capitella: In Chaetopterus (Glenner et al., 2004).
Haliotis: In Littorina (Gastropoda) (Glenner et al., 2004).
Mytilus: In Crassostrea (Bivalvia) (Glenner et al., 2004).
Serpula: “Groups such as Sabellariidae […] have evolved downstream-feeding without the aid of a metatroch” – (Rouse, 2000)In Chaetopterus (Glenner et al., 2004).
Sipunculus: “Taxa such as Sipuncula […] have a metatroch and do not have downstream larval-feeding” – Rouse (2000).
Terebratulina: Following closest relative in Glenner et al. (2004).

[239] Ciliary bands: Upstream

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 239: Larva: Cilia: Ciliary bands: Upstream

0: Absent1: Present?: AmbiguousNeomorphic character.

Upstream-collecting ciliary bands with single cilia on monociliated cells. Character 32 in Glenner et al. (2004).

Serpula, Capitella: In Chaetopterus (Glenner et al., 2004).
Haliotis: In Littorina (Gastropoda) (Glenner et al., 2004).
Mytilus: In Crassostrea (Bivalvia) (Glenner et al., 2004).
Terebratulina: Following closest relative in Glenner et al. (2004).

[240] Adoral ciliary band

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 240: Larva: Cilia: Adoral ciliary band

0: Absent1: Present?: AmbiguousNeomorphic character.

Characters 1.50, 2.66 and 4.68 in von Salvini-Plawen & Steiner (1996); 2 in Vinther et al. (2008). See also characters 39 in Haszprunar (1996) and 153 in Giribet & Wheeler (2002).

Haliotis: De Viçose et al. (2007).

[241] Nerve ring underlying ciliated larval swimming organ

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 241: Larva: Cilia: Nerve ring underlying ciliated larval swimming organ

0: Absent1: Present?: AmbiguousNeomorphic character.

Following Wanninger (2009).

Amathia: Nodular nerve ring underlies corona (Reed & Cloney, 1982).
Flustra: Present, following schematic in Gruhl & Schwaha (2016).

4.25 Ciliary ultrastructure

[242] Accessory centriole

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 242: Ciliary ultrastructure: Accessory centriole

0: Absent1: Present?: AmbiguousNeomorphic character.

After Lundin, Schander, & Todt (2009).

Serpula: Present in certain annelids; not verified in Serpula.
Terebratulina: Present (Lüter, 1995).

[243] Aggregation of granules below basal plate

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 243: Ciliary ultrastructure: Aggregation of granules below basal plate

0: Absent1: Present?: AmbiguousNeomorphic character.

After Lundin et al. (2009).

Serpula: Following Harmothoe (Holborow, Laverack, & Barber, 1969).

[244] Radiating tubular fibres

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 244: Ciliary ultrastructure: Basal foot: Radiating tubular fibres

0: Absent1: Present?: AmbiguousNeomorphic character.

After Lundin et al. (2009). Fibres radiate from the distal end of the basal foot of the cilia in certain taxa.

Amathia: Reed & Cloney (1982).
Serpula: Basal foot in Magelona is connected to cytoplasmic microtubules (Bartolomaeus, 1995).

[245] Basal plate

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[2]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 245: Ciliary ultrastructure: Basal plate

1: Thin2: Blurry?: AmbiguousTransformational character.

After Lundin et al. (2009). Also termed “dense plate”.

Amathia: Reed & Cloney (1982).
Serpula: Broad and ‘blurry’ in Magelona (Bartolomaeus, 1995).
Terebratulina: Thin to thick, but not blurry (Lüter, 1995).

[246] Brushborder of microvilli

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 246: Ciliary ultrastructure: Brushborder of microvilli

0: Absent1: Present?: AmbiguousNeomorphic character.

After Lundin et al. (2009); coded following Smith (2012a).

Amathia: Present (Reed & Cloney, 1982).
Terebratulina: Absent (Lüter, 1995).

[247] Centriolar triplet derivative in basal body

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[2]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 247: Ciliary ultrastructure: Centriolar triplet derivative in basal body

1: 9 + 2 pattern2: 9 + 3 pattern?: AmbiguousTransformational character.

After Lundin et al. (2009).

Amathia: Reed & Cloney (1982).
Serpula: Following Enchytraeus (Reger, 1967), Magelona (Bartolomaeus, 1995) and Harmothoe (Holborow et al., 1969).

[248] Ciliary necklace with connecting strands

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 248: Ciliary ultrastructure: Ciliary necklace with connecting strands

0: Absent1: Present?: AmbiguousNeomorphic character.

After Lundin et al. (2009).
The ciliary necklace is defined by Gilula & Satir (1972) as “Well-defined rows or strands of membrane particles that encircle the ciliary shaft”. It occurs immediately below the basal plate, and comprises three beaded circles of on the circumference of the cilia membrane.

Amathia: Reed & Cloney (1982).
Serpula: Not evident in Enchytraeus (Reger, 1967), Magelona (Bartolomaeus, 1995) or Harmothoe (Holborow et al., 1969).
Terebratulina: (Lüter, 1995).

[249] Monociliate epidermal cells

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 249: Ciliary ultrastructure: Monociliate epidermal cells

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 4 in Parry & Caron (2019). Coded as present if compound cilia comprise multiple monociliate cells, even if monociliate cells do not occur individually.

Serpula, Capitella, Mopalia, Tonicella, Leptochiton, Haliotis: Following coding for closest relative in Parry & Caron (2019).
Novocrania, Lingula, Odontogriphus omalus, Wiwaxia corrugata, Halkieria evangelista: Following coding in Parry & Caron (2019).

[250] Presence

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[1]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 250: Ciliary ultrastructure: Compound cilia: Presence

0: Absent1: Present?: AmbiguousNeomorphic character.

After Lundin et al. (2009). Compound cilia are motile structures composed of 10–100 regular cilia used in locomotion or feeding.

Amathia: Reed & Cloney (1982).
Capitella: Following closest relative in Glenner et al. (2004).
Serpula: Nielsen (1987).

[251] Origin

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[2]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[2]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[-]Flustra[-]Amathia[2]Loxosomella[?]Namacalathus[-]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 251: Ciliary ultrastructure: Compound cilia: Origin

1: Several monociliate cells2: On multiciliated cell?: Ambiguous-: InapplicableTransformational character.

Character 14 in Glenner et al. (2004). Compound cilia can be produced by the aggregation of cilia from multiple monociliate cells, or from a single cell bearing multiple cilia (Nielsen, 1987).

Capitella, Mytilus, Haliotis: Following closest relative in Glenner et al. (2004).
Terebratulina: “The coelothelial cells of the metacoel are monociliated”; “even some epithelial muscle cells are monociliated” – Lüter (1995).

[252] Glycocalyx ultrastructure

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 252: Ciliary ultrastructure: Glycocalyx ultrastructure

1: Homogeneous2: Layered?: AmbiguousTransformational character.

After Lundin et al. (2009).

Amathia: Reed & Cloney (1982).
Terebratulina: Homogeneous (Lüter, 1995).

[253] Branched

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 253: Ciliary ultrastructure: Microvilli on epidermal surface: Branched

0: Unbranched1: Branched?: AmbiguousNeomorphic character.

After Lundin et al. (2009).

Amathia: Reed & Cloney (1982).
Terebratulina: (Lüter, 1995).

[254] Length

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 254: Ciliary ultrastructure: Vertical ciliary rootlet: Length

1: Short2: Long?: AmbiguousTransformational character.

After Lundin et al. (2009). The vertical ciliary rootlet is also termed the posterior rootlet.

Amathia: Reed & Cloney (1982).
Loxosomella: Details of ciliary ultrastructure are illustrated in Nielsen & Rostgaard (1976).
Terebratulina: Long (Lüter, 1995).

[255] Shape

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[2]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 255: Ciliary ultrastructure: Vertical ciliary rootlet: Shape

1: Conical2: FlatTransformational character.

After Lundin et al. (2009). The vertical ciliary rootlet is also termed the posterior rootlet.

Amathia: Reed & Cloney (1982).
Serpula: Conical in Enchytraeus (Reger, 1967) and Magelona (Bartolomaeus, 1995).
Terebratulina: Conical: tapering to a point (Lüter, 1995).

[256] Presence

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[0]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 256: Ciliary ultrastructure: Secondary ciliary rootlet: Presence

0: Absent1: Present?: AmbiguousNeomorphic character.

After Lundin et al. (2009). The secondary ciliary rootlet is also termed the anterior ciliary rootlet.

Amathia: Reed & Cloney (1982).
Wirenia, Chaetoderma, Mopalia, Leptochiton: Lundin & Schander (2001).
Mytilus, Haliotis: Absent in class in adult stage (Lundin & Schander, 2001).
Neopilina: According to unpublished observation in Lundin & Schander (2001).

[257] Length

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[-]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[-]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 257: Ciliary ultrastructure: Secondary ciliary rootlet: Length

1: Short2: Long?: Ambiguous-: InapplicableTransformational character.

After Lundin et al. (2009). The secondary ciliary rootlet is also termed the anterior ciliary rootlet.

Amathia: Reed & Cloney (1982).
Serpula: Short in Enchytraeus (Reger, 1967), Magelona (Bartolomaeus, 1995) and Harmothoe (Holborow et al., 1969).
Terebratulina: “Very small” – Lüter (1995).

[258] Shape

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[2]Tonicella[-]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[-]Neopilina[?]Conocardium elongatum[-]Haliotis[?]Pojetaia runnegari[-]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[-]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 258: Ciliary ultrastructure: Secondary ciliary rootlet: Shape

1: Conical2: Flat?: Ambiguous-: InapplicableTransformational character.

After Lundin et al. (2009). The secondary ciliary rootlet is also termed the anterior ciliary rootlet.

Amathia: Reed & Cloney (1982).
Serpula: Conical in Magelona (Bartolomaeus, 1995).
Terebratulina: Too small to evaluate.

4.26 Nephridia

[259] Podocytes

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 259: Nephridia: Podocytes

0: Absent1: Present?: AmbiguousNeomorphic character.

See characters 21 and 28 in Haszprunar (2000); 1.12 in A. H. Scheltema (1993).

Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina: “In Brachiopoda, podocytes have never been observed” – Lüter (1995).
Phoronis: Present (Storch & Herrmann, 1978).
Serpula: Present in serpulids (Bartolomaeus & Quast, 2005).

[260] Rhogocytes

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 260: Nephridia: Rhogocytes

0: Absent1: PresentNeomorphic character.

Pore cells. Character 20 in Haszprunar (2000).

[261] Serve as excretory organs

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 261: Nephridia: Serve as excretory organs

0: No1: YesNeomorphic character.

See character 4.46 in von Salvini-Plawen & Steiner (1996).

Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina: “The excretory function of the metanephridia in Brachiopoda must be questioned” – Lüter (1995).

[262] Serially repeated

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[2]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 262: Nephridia: Serially repeated

1: Not repeated2: Serially repeated?: AmbiguousTransformational character.

Character 5 in Parry & Caron (2019).

Novocrania, Lingula, Sipunculus, Serpula, Capitella, Mopalia, Tonicella, Leptochiton, Haliotis: Following closest relative in Parry & Caron (2019).

[263] Protonephridia

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 263: Nephridia: Protonephridia

0: Absent1: Present?: AmbiguousNeomorphic character.

Also termed cyrtocytes. Character 21 in Grobe (2007); 1.47 in von Salvini-Plawen & Steiner (1996); 138 in Rouse (1999); 20 in Haszprunar (1996); 90 in Glenner et al. (2004).

[264] Metanephridia

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 264: Nephridia: Metanephridia

0: Absent1: PresentNeomorphic character.

See characters 35 in Rouse (1999); 28 in Haszprunar (2000); 93 in Glenner et al. (2004); 1.47 in von Salvini-Plawen & Steiner (1996); 21 in Grobe (2007); 138 in Rouse (1999); 20 in Haszprunar (1996).

Amathia: Following closest relative in Glenner et al. (2004).

4.27 Cuticle

[265] Layers

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 265: Cuticle: Layers

0: Simple (i.e. glycocalyx)1: Distinct epicuticle and endocuticle?: AmbiguousNeomorphic character.

Character 1 in Haszprunar (1996).

[266] Composition

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[1]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 266: Cuticle: Composition

1: Chitinous2: CollagenTransformational character.

Character 2 in Haszprunar & Wanninger (2008).

Flustra, Amathia: Collagenous (Schopf & Manheim, 1967), though chitin is associated with the exoskeleton (Hunt, 1972).
Tonicella, Dentalium: Haszprunar & Wanninger (2008).
Lingula, Pelagodiscus atlanticus, Terebratulina: The brachiopod pedicle has a chitinous cuticle (MacKay & Hewitt, 1978; Williams et al., 1997), but the tentacles are associated with collagen (Williams et al., 1997); marked as polymorphic.
Loxosomella: Absent (Haszprunar & Wanninger, 2008). Chitin is occasionally present in certain species, perhaps in regions where rigidity is necessary (Borisanova, Yushin, Malakhov, & Temereva, 2015).
Novocrania: No (chitinous) pedicle, so only collagenous cuticle present (Williams et al., 1997).
Phoronis: Collagen fibres in tentacle cuticle (Bartolomaeus, 2001); chitin only present in tubes (Jeuniaux, 1971).
Sipunculus: Collagenous (Goffinet, Voss-Foucart, & Barzin, 1978).

[267] Fibrous layer with thick fibrils

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 267: Cuticle: Fibrous layer with thick fibrils

0: Absent1: Present?: AmbiguousNeomorphic character.

After Borisanova et al. (2015).

Loxosomella, Flustra, Amathia, Serpula, Tonicella, Dentalium: Following table 2 in Borisanova et al. (2015).
Lingula: Pedicle cuticle entirely homogeneous (Williams et al., 1997).
Pelagodiscus atlanticus: Microvilli in otherwise homogeneous epidermis (Williams et al., 1997).
Phoronis: Outer layer seemingly fibrous (Bereiter-Hahn, Matoltsy, & Slyvia Richards, 1984).
Sipunculus: Fibrous collagen only (Bereiter-Hahn et al., 1984).
Terebratulina: Not evident in Notosaria (Bereiter-Hahn et al., 1984; Williams et al., 1997).

[268] Homogeneous layer

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 268: Cuticle: Homogeneous layer

0: Absent1: Present?: AmbiguousNeomorphic character.

After Borisanova et al. (2015).

Loxosomella, Flustra, Amathia, Serpula, Tonicella, Dentalium: Following table 2 in Borisanova et al. (2015).
Lingula: Pedicle cuticle entirely homogeneous (Williams et al., 1997).
Pelagodiscus atlanticus: Microvilli in otherwise homogeneous epidermis (Williams et al., 1997).
Phoronis: Not evident (Bereiter-Hahn et al., 1984).
Sipunculus: Fibrous collagen only (Bereiter-Hahn et al., 1984).
Terebratulina: Cuticle is homogeneous in Notosaria (Bereiter-Hahn et al., 1984; Williams et al., 1997).

[269] Resilience

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 269: Cuticle: Resilience

0: Labile1: Robust?: Neomorphic character.

Character 1 in Haszprunar (2000).

[270] Microvilli

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 270: Cuticle: Microvilli

0: Absent1: Microvilli present in the cuticle?: AmbiguousNeomorphic character.

After Borisanova et al. (2015).

Loxosomella, Flustra, Amathia, Serpula, Tonicella, Dentalium: Following table 2 in Borisanova et al. (2015).
Pelagodiscus atlanticus: Microvillios inner epithelium in Discina (Williams et al., 1997).
Phoronis: Present on outer epithelium (Bereiter-Hahn et al., 1984).
Sipunculus: Fibrous collagen only (Bereiter-Hahn et al., 1984).

4.28 Muscles

[271] Longitudinal muscle bands

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 271: Muscles: Longitudinal muscle bands

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 127 in Parry & Caron (2019).

Novocrania, Lingula, Sipunculus, Canadia spinosa, Serpula, Capitella, Mopalia, Tonicella, Leptochiton, Haliotis, Odontogriphus omalus, Wiwaxia corrugata, Halkieria evangelista: Following closest relative in Parry & Caron (2019).

[272] Circular muscles

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 272: Muscles: Circular muscles

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 128 in Parry & Caron (2019).

Novocrania, Lingula, Sipunculus, Canadia spinosa, Serpula, Capitella, Mopalia, Tonicella, Leptochiton, Haliotis, Odontogriphus omalus, Wiwaxia corrugata, Halkieria evangelista: Following closest relative in Parry & Caron (2019).

[273] Cytology

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[2]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[3]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[3]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[3]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[3]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 273: Muscles: Cytology

1: Smooth2: Obliquely striated3: Smooth on abfrontal face; striated on frontal face?: AmbiguousTransformational character.

Character 19 in Haszprunar (1996); see also character 13 in Haszprunar (2000).

Flustra, Amathia: In Bryozoa, myofibrils are “all striated” (Pardos et al., 1991).
Novocrania, Lingula, Pelagodiscus atlanticus, Terebratulina: In brachiopods, myofibrils “are striated on the frontal face and smooth on the abfrontal face” (Pardos et al., 1991).
Phoronis: “In P. australis […] all the myofibrils belong to the smooth type” – Pardos et al. (1991).

[274] Histology

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 274: Muscles: Histology

0: Fibre-type1: Epithelially organized?: Neomorphic character.

See character 18 in Haszprunar (1996).

4.29 Glands: Paired pharyngeal diverticulae [275]

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 275: Glands: Paired pharyngeal diverticulae

0: Absent1: PresentNeomorphic character.

4.30 Nervous system

[276] Orthogonal

Character adds 4 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 276: Nervous system: Orthogonal

0: Not orthogonal1: Orthogonal?: AmbiguousNeomorphic character.

Character 14 in Haszprunar (1996). Paired longitudinal nerve cords regularly interconnected by transversal commissures to form a rectangular pattern.

Amathia: Temereva & Kosevich (2016).
Canadia spinosa, Capitella: Parry & Caron (2019).
Haliotis: Pedal cords with regular transverse connections (Barlow & Truman, 1992).
Mopalia, Tonicella, Leptochiton: Parry & Caron (2019) code chitons as lacking serial repetition in the nervous system, despite the orthogonal arrangement of nervous tissue.

[277] Glial system

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 277: Nervous system: Glial system

0: Absent1: PresentNeomorphic character.

Character 16 in Haszprunar (1996). The Gliointerstitial system interconnects the nervous and muscle systems.

Phoronis: Glial cells are “abundant” (Temereva, 2016).

[278] Dorso-terminal sense organ

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[?]Sipunculus[1]Wirenia[1]Chaetoderma[?]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 278: Nervous system: Dorso-terminal sense organ

0: Absent1: Present?: AmbiguousNeomorphic character.

Corresponds to the molluscan osphradium; see (???), character 30; Ponder & Lindberg (1997), character 101; Giribet & Wheeler (2002) character 143; Haszprunar (2000) character 56; (???) character 49; (???) character 101.

[279] Statocysts

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[0]Serpula[?]Capitella[?]Canadia spinosa[?]Sipunculus[0]Wirenia[0]Chaetoderma[?]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[0]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 279: Nervous system: Statocysts

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 1.33 in (???); 44 in (???); 99 in Ponder & Lindberg (1997); 55 in Haszprunar (2000).

[280] Nuchal organs

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 280: Nervous system: Nuchal organs

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 147 in L. A. Parry et al. (2016), 158 in Parry & Caron (2019).
Nuchal organs are chemosensory organs present in almost all polychaetes, and absent in clitellates. They occur as a dorsal pair of ciliated areas on the posterior prostomium (???). (???) points to a number of differences between the nuchal organs of sipunculans and polychaetes, whilst acknowledging the existence of some similarities; (???) acknowledge that the case is not closed. We agree that homology between the nuchal organs of sipunculans and annelids is uncertain, but code the structures in a single transformation series to allow the analysis to test the hypothesis of homology.

[281] Buccal nerve ring

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[?]Capitella[1]Canadia spinosa[?]Sipunculus[1]Wirenia[0]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 281: Nervous system: Buccal nerve ring

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 7b in Haszprunar & Wanninger (2008), following Wanninger et al. (2007). Also termed an oral or circumoral nerve ring.

Amathia: Temereva & Kosevich (2016).
Sipunculus, Canadia spinosa, Serpula: Coded as present in Parry & Caron (2019).
Haliotis: Present (Barlow & Truman, 1992).
Loxosomella: A potential synapomorphy uniting Mollusca and Kamptozoa (???; Wanninger et al., 2007).

[282] Anterior nerve loop

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 282: Nervous system: Anterior nerve loop

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 7c in Haszprunar & Wanninger (2008), following Wanninger et al. (2007). An anterior, pre-oral nerve loop is present in molluscs, Loxosomella and certain annelids (Wanninger et al., 2007).

Amathia: Temereva & Kosevich (2016).
Haliotis: Loop joins cerebral ganglia anterior to buccal loop (Barlow & Truman, 1992).
Mopalia, Leptochiton: Present in polyplacophorans (Wanninger et al., 2007).

[283] Suprarectal commissure

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[?]Capitella[?]Canadia spinosa[?]Sipunculus[1]Wirenia[1]Chaetoderma[?]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 283: Nervous system: Suprarectal commissure

0: Absent1: Present?: AmbiguousNeomorphic character.

Also termed suprarectal loop; see (???) character 28; A. H. Scheltema (1993) character 21; Waller (1998) character 2e.

Haliotis: Barlow & Truman (1992).

[284] Ganglionated

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[?]Capitella[?]Canadia spinosa[?]Sipunculus[1]Wirenia[1]Chaetoderma[?]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[0]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[0]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[?]Amathia[0]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 284: Nervous system: Suprarectal commissure: Ganglionated

0: Ganglia absent1: Strongly ganglionated?: AmbiguousNeomorphic character.

[285] Formation of ganglia

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[3]Flustra[3]Amathia[1]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[2]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[2]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 285: Nervous system: Formation of ganglia

1: From cerebral region2: In situ3: Invagination of epithelium?: AmbiguousTransformational character.

Character 1.22 in von Salvini-Plawen & Steiner (1996).

Flustra, Amathia: “The cerebral ganglion in all bryozoans is formed as an invagination of a portion
of epithelium” – Temereva & Kosevich (2016).

[286] Presence

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[1]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 286: Nervous system: Cerebral ganglia: Presence

0: Absent1: Present?: AmbiguousNeomorphic character.

After character 13 in Haszprunar (1996).

Amathia: Temereva & Kosevich (2016).
Haliotis: Barlow & Truman (1992).
Phoronis: We treat the dorsal ganglion, which is formed by two ends of the tentacular nerve ring (Temereva, 2016), as cerebral.

[287] Fused

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[2]Serpula[?]Capitella[?]Canadia spinosa[2]Sipunculus[2]Wirenia[2]Chaetoderma[2]Tonicella[2]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[2]Amathia[2]Loxosomella[?]Namacalathus[2]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[-]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[-]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 287: Nervous system: Cerebral gangila: Fused

1: Pair of distinct ganglia2: Single ganglion, or fused ganglia?: Ambiguous-: InapplicableTransformational character.

After character 13 in Haszprunar (1996).

Amathia: Fused (Temereva & Kosevich, 2016).
Haliotis: Barlow & Truman (1992).

[288] Transverse commissures

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 288: Nervous system: Cerebral ganglia: Transverse commissures

0: Absent1: Present?: AmbiguousNeomorphic character.

Character 205 in Parry & Caron (2019), who write “a brain with four transverse commissures is present in numerous families of polychaetes and two commissures are present in Sipuncula”.

Capitella: The brain of Notomastus (Capitellidae) comprises two brain lobes joined by a commissure (???).
Mopalia, Tonicella, Leptochiton, Haliotis: Following closest relative in Parry & Caron (2019).

[289] Serially repeated ganglia

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 289: Nervous system: Serially repeated ganglia

0: Absent1: PresentNeomorphic character.

Character 212 in Parry & Caron (2019).

Novocrania, Lingula, Sipunculus, Canadia spinosa, Serpula, Capitella, Mopalia, Tonicella, Leptochiton, Haliotis, Odontogriphus omalus, Wiwaxia corrugata, Halkieria evangelista: Following closest relative in Parry & Caron (2019).

[290] Serially repeated segmental nerves

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[0]Sipunculus[?]Wirenia[?]Chaetoderma[0]Tonicella[0]Mopalia[?]Polysacos vickersianum[0]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 290: Nervous system: Serially repeated segmental nerves

0: Absent1: PresentNeomorphic character.

Character 213 in Parry & Caron (2019).

Novocrania, Lingula, Sipunculus, Canadia spinosa, Serpula, Capitella, Mopalia, Tonicella, Leptochiton, Haliotis: Following closest relative in Parry & Caron (2019).

[291] Nerve cords

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[2]Wirenia[2]Chaetoderma[2]Tonicella[2]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[2]Haliotis[?]Pojetaia runnegari[?]Mytilus[2]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[1]Flustra[1]Amathia[2]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[1]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[1]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 291: Nervous system: Nerve cords

1: Ventral nerve cords only2: Tetraneury: one pair of ventral and one pair of lateral nerve cordsTransformational character.

See character 7 in Haszprunar & Wanninger (2008).

Amathia: Temereva & Kosevich (2016).

[292] Medulla

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[?]Capitella[?]Canadia spinosa[?]Sipunculus[2]Wirenia[2]Chaetoderma[?]Tonicella[2]Mopalia[?]Polysacos vickersianum[2]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[2]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 292: Nervous system: Nerve cords: Medulla

1: Not medullary2: MedullaryTransformational character.

Medullary nerve cord are built by a core neuropil and covered by neuronal somata (???).

[293] Ventral longitudinal nerves

Character adds 3 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[?]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[1]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[0]Flustra[0]Amathia[1]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[0]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[0]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[0]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[0]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 293: Nervous system: Ventral longitudinal nerves

0: Separate1: Paired or secondarily fused?: AmbiguousNeomorphic character.

Character 80 in Glenner et al. (2004); see also character 6 in Vinther et al. (2008).

Amathia: Temereva & Kosevich (2016).
Haliotis: Linked but distinct pedal cords (Barlow & Truman, 1992).

[294] Ventral cord location

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[2]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[2]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 294: Nervous system: Ventral cord location

1: Subepidermal2: Intraepidermal?: AmbiguousTransformational character.

Character 222 in Parry & Caron (2019).

Novocrania, Lingula, Sipunculus, Canadia spinosa, Serpula, Capitella, Mopalia, Tonicella, Leptochiton, Haliotis: Following coding in Parry & Caron (2019), who follow (???).

[295] Ventral cord commissures

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[2]Capitella[2]Canadia spinosa[1]Sipunculus[?]Wirenia[?]Chaetoderma[1]Tonicella[1]Mopalia[?]Polysacos vickersianum[1]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[?]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[-]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[-]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 295: Nervous system: Ventral cord commissures

1: Not segmentally arranged2: Segmentally arranged-: InapplicableTransformational character.

Character 223 in Parry & Caron (2019).

Novocrania, Lingula, Sipunculus, Canadia spinosa, Serpula, Capitella, Mopalia, Tonicella, Leptochiton, Haliotis: Coded following Parry & Caron (2019), who follow (???).

4.31 MicroRNA

[296] Brachiopod candidate 1

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[?]Capitella[?]Canadia spinosa[?]Sipunculus[?]Wirenia[?]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[0]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 296: MicroRNA: Brachiopod candidate 1

0: Absent1: Present?: AmbiguousNeomorphic character.

BC1 in 35.

Novocrania, Lingula, Phoronis, Mytilus: 35.

[297] mir-36

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 297: MicroRNA: mir-36

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[298] mir-76

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[1]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 298: MicroRNA: mir-76

0: Absent1: PresentNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[299] mir-124

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 299: MicroRNA: mir-124

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[300] mir-190

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 300: MicroRNA: mir-190

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[301] mir-219

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 301: MicroRNA: mir-219

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[302] mir-242

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 302: MicroRNA: mir-242

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[303] mir-278

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[1]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[1]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 303: MicroRNA: mir-278

0: Absent1: PresentNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[304] mir-1984

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 304: MicroRNA: mir-1984

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[305] mir-1985

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 305: MicroRNA: mir-1985

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[306] mir-1986

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 306: MicroRNA: mir-1986

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[307] mir-1987

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 307: MicroRNA: mir-1987

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[308] mir-1988

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 308: MicroRNA: mir-1988

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[309] mir-1989

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 309: MicroRNA: mir-1989

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[310] mir-1990

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[1]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 310: MicroRNA: mir-1990

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[311] mir-1991

Character adds 2 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[1]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 311: MicroRNA: mir-1991

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[312] mir-1994

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[1]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 312: MicroRNA: mir-1994

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[313] mir-1995

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 313: MicroRNA: mir-1995

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[314] mir-1996

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 314: MicroRNA: mir-1996

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[315] mir-1997

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 315: MicroRNA: mir-1997

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[316] mir-1998

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 316: MicroRNA: mir-1998

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[317] mir-1999

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 317: MicroRNA: mir-1999

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[318] mir-2000

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 318: MicroRNA: mir-2000

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[319] mir-2001

Character adds 0 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[1]Serpula[1]Capitella[1]Canadia spinosa[1]Sipunculus[1]Wirenia[1]Chaetoderma[1]Tonicella[1]Mopalia[1]Polysacos vickersianum[1]Leptochiton[1]Glaphurochiton carbonarius[1]Calvapilosa kroegeri[1]Phthipodochiton thraivensis[1]Wiwaxia corrugata[1]Odontogriphus omalus[1]Neopilina[1]Conocardium elongatum[1]Haliotis[1]Pojetaia runnegari[1]Mytilus[1]Dentalium[1]Pelagiella[1]Siphogonuchites multa[1]Acaenoplax hayae[1]Kulindroplax perissokomos[1]Paramicrocornus[1]Haplophrentis carinatus[1]Cupitheca holocyclata[1]Bactrotheca[1]Pedunculotheca diania[1]Halkieria evangelista[1]Flustra[1]Amathia[1]Loxosomella[1]Namacalathus[1]Phoronis[1]Dailyatia[1]Cotyledion tylodes[1]Eccentrotheca[1]Orthrozanclus[1]Micromitra[1]Askepasma toddense[1]Coolinia pecten[1]Antigonambonites planus[1]Yuganotheca elegans[1]Tomteluva perturbata[1]Salanygolina[1]Terebratulina[1]Orthis[1]Alisina[1]Longtancunella chengjiangensis[1]Glyptoria[1]Nisusia sulcata[1]Kutorgina chengjiangensis[1]Novocrania[1]Craniops[1]Ussunia[1]Heliomedusa orienta[1]Gasconsia[1]Pelagodiscus atlanticus[1]Mickwitzia muralensis[1]Mummpikia nuda[1]Botsfordia[1]Eoobolus[1]Lingulellotreta malongensis[1]Lingula[1]Siphonobolus priscus[1]Acanthotretella spinosa[1]Lingulosacculus[1]Clupeafumosus socialis[1]Micrina[1]Paterimitra[1]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 319: MicroRNA: mir-2001

1: PresentNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[320] mir-2685

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 320: MicroRNA: mir-2685

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[321] mir-2686

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 321: MicroRNA: mir-2686

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[322] mir-2687

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 322: MicroRNA: mir-2687

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[323] mir-2688

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 323: MicroRNA: mir-2688

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[324] mir-2689

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 324: MicroRNA: mir-2689

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[325] mir-2690

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 325: MicroRNA: mir-2690

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[326] mir-2691

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[1]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 326: MicroRNA: mir-2691

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[327] mir-2693

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 327: MicroRNA: mir-2693

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[328] mir-2693

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[1]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[0]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[0]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 328: MicroRNA: mir-2693

0: Absent1: Present?: AmbiguousNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[329] mir-2722

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[0]Capitella[?]Canadia spinosa[0]Sipunculus[?]Wirenia[1]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[1]Haliotis[?]Pojetaia runnegari[?]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[?]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[?]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 329: MicroRNA: mir-2722

0: Absent1: PresentNeomorphic character.

Phoronis, Capitella, Chaetoderma, Haliotis: 34.
Sipunculus: Following Phascolion (???).

[330] mir-5045

Character adds 1 to tree score0 additional regionsAn optimal tree under:Implied weights, k = 4.5IW, k = 10.5IW, k = 24Equal weights[?]Serpula[?]Capitella[?]Canadia spinosa[?]Sipunculus[?]Wirenia[?]Chaetoderma[?]Tonicella[?]Mopalia[?]Polysacos vickersianum[?]Leptochiton[?]Glaphurochiton carbonarius[?]Calvapilosa kroegeri[?]Phthipodochiton thraivensis[?]Wiwaxia corrugata[?]Odontogriphus omalus[?]Neopilina[?]Conocardium elongatum[?]Haliotis[?]Pojetaia runnegari[0]Mytilus[?]Dentalium[?]Pelagiella[?]Siphogonuchites multa[?]Acaenoplax hayae[?]Kulindroplax perissokomos[?]Paramicrocornus[?]Haplophrentis carinatus[?]Cupitheca holocyclata[?]Bactrotheca[?]Pedunculotheca diania[?]Halkieria evangelista[?]Flustra[?]Amathia[?]Loxosomella[?]Namacalathus[0]Phoronis[?]Dailyatia[?]Cotyledion tylodes[?]Eccentrotheca[?]Orthrozanclus[?]Micromitra[?]Askepasma toddense[?]Coolinia pecten[?]Antigonambonites planus[?]Yuganotheca elegans[?]Tomteluva perturbata[?]Salanygolina[?]Terebratulina[?]Orthis[?]Alisina[?]Longtancunella chengjiangensis[?]Glyptoria[?]Nisusia sulcata[?]Kutorgina chengjiangensis[1]Novocrania[?]Craniops[?]Ussunia[?]Heliomedusa orienta[?]Gasconsia[?]Pelagodiscus atlanticus[?]Mickwitzia muralensis[?]Mummpikia nuda[?]Botsfordia[?]Eoobolus[?]Lingulellotreta malongensis[1]Lingula[?]Siphonobolus priscus[?]Acanthotretella spinosa[?]Lingulosacculus[?]Clupeafumosus socialis[?]Micrina[?]Paterimitra[?]Yilingia spiciformis0.060.061110.060.060.060.0610.3310.060.060.060.060.060.0611110.170.110.170.170.940.940.060.140.92110.060.060.610.030.0610.22110.060.060.060.060.50.810.060.060.060.060.4710.060.360.810.6110.060.060.940.060.060.0611110.440.06

Character 330: MicroRNA: mir-5045

0: Absent1: Present?: AmbiguousNeomorphic character.

Novocrania, Lingula, Phoronis, Mytilus: 35.
Tree number:
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References

Adrianov, A. V., Maiorova, A. S., & Malakhov, V. V. (2011). Embryonic and larval development of the peanut worm Phascolosoma agassizii (Keferstein 1867) from the Sea of Japan (Sipuncula: Phascolosomatidea). Invertebrate Reproduction and Development, 55(1), 22–29. doi: 10.1080/07924259.2010.548638

Adrianov, A. V., Malakhov, V. V., & Maiorova, A. S. (2006). Development of the tentacular apparatus in sipunculans (Sipuncula): I. Thysanocardia nigra (Ikeda, 1904) and Themiste pyroides (Chamberlin, 1920). Journal of Morphology, 267(5), 569–583. doi: 10.1002/jmor.10423

Altenburger, A., & Wanninger, A. (2010). Neuromuscular development in Novocrania anomala: evidence for the presence of serotonin and a spiralian-like apical organ in lecithotrophic brachiopod larvae. Evolution and Development, 12(1), 16–24. doi: 10.1111/j.1525-142X.2009.00387.x

Altenburger, A., Wanninger, A., & Holmer, L. E. (2013). Metamorphosis in Craniiformea revisited: Novocrania anomala shows delayed development of the ventral valve. Zoomorphology, 132(4), 379–387. doi: 10.1007/s00435-013-0194-3

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